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ANNALS
OF
The Entomological Society of America
VOU ME ski, i909
EDITORIAL BOARD
H. COMSTOCK,
L. O. HOWARD,
TeisevA ae Ne, Yi
WASHINGTON, D. C.
- J. Ss. BETHUNE, W. M. WHEELER,
GUELPH, ONTARIO, CANADA. Boston, MASss.
Py Pay CAT VER.
PHILADELPHIA, Pa.
. L. KELLOGG, J. W. FOLSOM,
STANFORD UNIVv., CAL. URBANA, ILLS.
Boston, MASss.
HERBERT OSBORN, Managing Editor,
COLUMBUS, OHIO.
PUBLISHED QUARTERLY BY THE SOCIETY
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CONTENTS OF VOLUME II.
PAGE
Proceedings of the Entomological Society of America........... 1
Resolutions on the Death of Dr. Wm. H. Ashmead............. 6
Resolutions on the Death of Dr. James Fletcher...........:... ff
Report of the'Committeeron, Nomenclature.....2.....1..20:... 8
Comstock, J. H—A Note on the Habits of the Wall-bee Chali-
CO cd Omee etree cee ei ea nk Oe See Pe 8 cbsal water et gis mere Bhar va 9
PETRUNKEVITCH, A.—Contributions to Our Knowledge of the
Anatomysand: Relationships of Spiders. . o.......2..0vbs-- 11
GrRAULT, A. ARSENE—A Monographic Catalogue of the Mymarid
Genus Camptoptera Foerster, with Description of One New
Nosthmamentcany Norm.) 28.0. 224es wie Roe ote glee anaes 22
DAMS OrN |: ovudies on Aphididae ml: 0s. 202. cess. 30
Hitton, Witi1am A.—The Tracheal Supply in the Central
Nervous System of the Larva of Corydalis Cornuta......... 46
Netson, Jas. A.—Evolution and Adaptation in the Palpus of
Wil @uIkes (Sy SUG [Ss Sea a onl RR Be eae OU ee Nc 60
WesstTer, F. M.—Investigations of Toxoptera Graminum and
ISS: JERE eeENSS eK uae ae ee Oa ee AA es 67
Hayuurst, Paut—Observations on a Gall Aphid (Aphis
NGS ONGC Lo) et rane oe a en eer eg Be Sent hoe a eee 88
PatcH, Epirh M.—Homologies of the Wing Veins of the Aphi-
didae, Psyllidae, Aleurodidae, and Coccidae............... 101
Paine |AMES o:——Robpbertiies of the Genus Asilus.............. 136
CHAMBERLIN, RALPH V.—Some Records of North American
Geophilidae and Lithobiidae, with Description of New Species 175
Davis, JOHN J.—Two New Genera and Species of Aphididae.... 196
PouLton, Pror. E. B.—Mimicry in the Butterflies of North
[TGC AS OS ae Pe nae a tn 203
TOWNSEND, CHaAs. H. T.—Descriptions of New Genera and
SPeciessOlmnaeminiGACe rs. 6... Ase Pb asks aa Peloelsis body welels 243
CocKERELL, T. D. A.—Fossil Insects from Florissant........... PA
McGI.iivray, A. D.—A Synopsis of the North American Species
Ol OCOMOME IRI Cee p I ot fs bg.2 Cite oud ec wie s LR ao « 259
Rey era
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ANNALS
OF
The Entomological Society of America
Volume I] MAR GH, 1909 Number |
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF
AMERICA.
The fourth meeting of the Entomological Society of America
was held in Baltimore, December 30th and 31st, 1908, in affiliation
with the American Association for the Advancement of Science.
The meeting was called to order by the President, Dr. W. M.
Wheeler, in the Eastern High School, at 11 A. m., December 31st.
The minutes of the previous meeting were adopted as printed.
Dr. Fernald read the report of the committee on nomenclature.
Moved and carried that this report be received and printed and
discussed one year later, and that this should be the general
policy in dealing with these reports. This report is appended
to the minutes.
The Managing Editor made an informal report upon the con-
dition of the Annals. Moved and carried that the chair appoint
a committee on nominations. The chair appointed Messrs. C. W.
Johnson, C. L. Marlatt and H. T. Fernald. The President
announced the death during the year of Dr. W. H. Ashmead, an
Honorary Fellow, Dr. James Fletcher, a Fellow; and of three mem-
bers, C. Abbott Davis, Albert V. Taylor and Alexander Craw.
The President: announced that he had appointed Committees to
draw up suitable resolutions upon the death of Drs. Ashmead and
Fletcher. These resolutions were then read by the Secretary
and adopted as read. They are appended to these minutes. A
letter from Dr. L. O. Howard, concerning a request from Dr.
Carlos E. Porter for separata and an offer to specialists of material
for study, was read, and Mr. Webster made some explanations
thereto. The Secretary read a communication from W. C. Wood
suggesting action looking toward abolishing the tariff on insects.
Moved and carried, ‘‘That it is the sense of this Society that the
duty on insects is objectionable and should be abolished.” The
matter was referred to the Executive Committee with power.
Adjournment until 1:00 P. M.
2 Annals Entomological Society of America KViol,. Seis
At that time the meeting was called to order by the First
Vice President, Dr. J. B. Smith. The report of the nominating
committee was read, and in accordance therewith, the Secretary
was instructed to cast a ballot for the following officers, which was
done:
President, Dr. Henry Skinner.
First Vice President, Prof. Herbert Osborn.
Second Vice President, Dr. A. D. Hopkins.
Secretary—Treasurer, J. Chester Bradley.
Additional Members of the Executive Committee:
Prof. J. Hi. Comstock; Dr. John B. Smith,
Dr. W. M. Wheeler, neve bron GC. |. >. Bethune;
Mr. E. A. Schwarz, Prof. Lawrence Bruner.
Member of Committee on Nomenclature to serve three years,
Dr. E. P. Felt, to succeed himself.
There being no further business, papers were read as follows:
W.C. O’Kane. ‘‘Notes on a Lecanio-diaspid.”’
A. N. Caudell spoke briefly on a method that he had adopted
for preserving types. He uses Riker mounts for this purpose.
W.A. Hooker. ‘‘Contributions to our knowledge of the host
relations of ticks.’’ Discussion by Messrs. Skinner, Cooley, Vier-
eck, Bruner and Miss Mitchell.
The meeting then adjourned.
At 3 P. M. a joint session with Section F of the American Asso-
ciation for the Advancement of Science was held in the Eastern
High School. Dr. J. B. Smith presided over the meeting. The
following papers were read:
Mr. F. M. Webster. ‘‘Investigations of Toxoptera graminum
and its parasites.’’ Discussion by Prof. Washburn and Mr. Hay-
hurst.
A. Petrunkevitch. ‘‘On the muscular system of spiders’ legs.”’
(Published in this number of THE ANNALS.)
J. H. Comstock. ‘‘A note on the habits of the wall-bee, Chal-
icodoma.”’ (Published in full elsewhere in this volume of THE
ANNALS.)
J.A.WNelson. ‘Evolution and adaptation in the palpi of male
spiders.”” (To be published in full in THE AnNAts.) Discussion
by Drs. Hopkins and Petrunkevitch.
Professor Williston now took the chair while the following
paper was read:
1909] Proceedings 3
Jeb SMe es opecies,. varieties, races; etc: Discussion
was postponed until after the reading of the next.
H. Skinner. ‘‘What is a species?’’ Considerable discussion
was evoked by these two papers, Messrs. Williston, Fernald, Felt,
Viereck, Schwarz, Caudell and others participating.
Adjournment.
At 10 A. M. on December 31st, the meeting was called to order
by Dr. Wheeler, and the following papers read:
H. P. Severin. ‘‘Death-feigning by Zaitha fluminea.”’ Read
by Prof. Osborn.
C. R. Crosby. ‘‘Some habits of seed infesting Chalcis flies.”’
W. A. Riley. “‘The development of the scent-pockets of
Anosia.”’
W.A. Hilton. ‘‘Tracheal distribution in the nervous system
of larval Corydalis cornuta.”’
E. P. Felt. “Observations on the taxonomy of the Ceci-
domyiidae.”’
F. S. Sherman, Jr. ‘Recording and mapping the entomo-
logical fauna of a state.’’
D. W. Coquillett. “‘Rediscovery .of the Bibionid genus
Eupeitenus.”
In the absence of the authors, or for other reasons, the fol-
lowing papers were read by title only:
». A. Forbes, ‘‘Local relations of allied species.”’ A. Petrunke-
vitch, ‘“On some terms used in systematics.” E. A. Andrews,
“A colony of mound-building ants.”” W. E. Ritter, ‘‘The concep-
tion of unit-systems in biology.” C. T. Brues, ‘‘Some further
remarks on the systematic affinities of the Phoridae.” <A. A.
Girault, ““A monographic catalogue of the Myrmarid genus
Camptoptera Foerster, with description of one new North Ameri-
can form. |. ). Davis, “Studies on Aphididae II.”’ R. V-
Chamberlin, ‘‘A Newportia in Utah.”
The following report from the Executive Committee was
read and, with all its provisions and recommendations, adopted:
The Executive Committee announces the election of the following
Fellows: Samuel W. Williston, Theodore Dru Allison Cockerell, Eph-
raim Porter Felt, Elmer Darwin Ball, Alexander Dyer MacGillivray.
In June the following nine members were elected: W. O. Hart, 134
Carondelet St., New Orleans, La.; B. F. Berrenger, 3-4 Masonic Block,
Fostoria, O.; Prof. E. P. Durrant, Westerville, O.; Henrik Peder Lod-
ing, 911 Palmetto St., Mobile, Ala.; W. P. Flint, Office State Ento-
mologist, Urbana, Ill.; Charles N. Ainslie, Bureau Entomology, Wash-
ington, D. C.; Prof. A. F. Conradi, Clemson College, S. C.; H. H. Sever-
4 Annals Entomological Society of America (Mol. LL;
in, Columbus, O.; Walter Collins O’ Kane, Columbus, O. At this meet-
ing forty-two more have been elected as follows: Mrs. A. W. Smith,
15 E. Ave., Ithaca, N. Y.; Miss Edna Mosher, 801 Armistead Ave.,
Hampton, Va.; Miss Hortense Butler, Peterson, Ia.; Mr. Chas. H. Mer-
genthaler, 113 N. Main St., Fostoria, O.; Mr. C. H. Withington, Law-
rence, Kan.; Mr. W. Reiff, Bussey Institution, Forest Hills, Mass. ;
Mr. F. A. Johnston, 84 Pleasant St., Amherst, Mass.; Mr. D. B’ Kuhns,
Honolulu; Mr. J. N. Summers, Insectary, Amherst; Mr. J. A. Hyslop,
205 D. St., N. W., Washington, D. C.; Mr. E. G. Smyth, 1100 Va. Ave.,
S. W., Washington, D. C.; Mr. J. F. Zimmer, 111 W. 11th Ave., Colum-
bus, O.; Mr. R. H. Wolcott, Lincoln, Neb.; Mr. H. T. Osborn, 485 King
Ave., Columbus, O.; Mr. V. L. Wildermuth, Groveport, O.; Mr. W. H.
Goodwin, 39 Beall Ave., Wooster, O.; Mr. A. W. McCray, Bur. Entom.,
Washington, D. C.; Mr. H. C. Severin, 941 Grove St., Milwaukee, Wis. ;
Mr. B. M. Chatterjie, 37 Jaliapara Road, Bhowanipur, Calcutta; Mr.
‘ W. T. M. Forbes, 614 E. State St., Ithaca; Mr. J. P. Jensen, Forest
Home, Ithaca; Mr. W. Moore, Ithaca; Miss A. H. Morgan, Mt. Holyoke
College, S. Hadley, Mass.; Mr. A. Dutt, Coochbehar, Bengal; Mr. W.S.
Regan, Amherst; Mr. C. E. Hood, Box 208, Dallas, Tex.; Mr. W. A.
Hilton, 108 Brandon Place, Ithaca; Mr. W. J. Price, Blacksburg, Va. ;
Mr. Z. P. Metcalf, Raleigh, N. C.; Mr. W. P. Comstock, Newark, N. J.;
Mr. S. A. Rohwer, Boulder, Col.; Mr. O. Bryant, Cohasset, Mass.; Miss
S; Di McCann, 137 E. High St., Lexington, Kye; Mr .W. F. Turner;
Box O, Auburn, Ala.; Mr. C. J. Triggerson, 211 Hudson St., Ithaca;
Mr. P. H. Hertzog, Lewisburg, Pa.; Mr. R. W. Braucher, Bureau
Entom., Washington, D. C.; Mr. H. Dean, Bureau of Entom., Wash-
ington, D. C.; Mr. Nathan Banks, East Falls Church, Va.; Mr. C. P.
Smith, Logan, Utah; Mr. H. G. Cribs, Philadelphia; Mr. L. L. Scott,
Columbus, O.
The following suggestion from Prof. Cockerell, it is recommended,
be referred to the Committee on Nomenclature: ‘‘That the Entomo-
logical Society undertake to get out a list of all North American insects
to be used as a standard like the A. O. U. Code. Of course it would
take years and would have to come out in parts.”’
‘“Reverend Professor C. J.S. Bethune was elected a member of the
Editorial Board to fill the vacancy caused by the death of Dr. Fletcher.
‘““The Committee recommends the following amendment to the
Constitution, to be voted on at the next annual meeting:
‘‘Art. 5, Sec. 3 to omit the following: ‘All Officers, Sec-
retary-Treasurer excepted, and all additional members of the
Executive Committee, shall be chosen from the list of Fellows.
Provided, etc.; so that the section will read: ‘Art. V, Sec. 3.
Election of Officers. All officers shall be elected by ballot at
the annual meeting for a term of one year and shall be eligible
for re-election.’
‘The Committee recommends the adoption of the following by-law:
‘6. Any member may become a life member upon pay-
ment of $50.00 at one time, and shall be exempt from further
assessments. He shall receive during his life, one copy of
each issue of the ANNALS.
1909] Proceedings 5
‘The resignations of Messrs. G. W. Lucas, C. S. Brimley, W. F.
Works, T. W. Fyles, and Miss F. H. Goodfellow were accepted.
“The following is a summary of the financial condition of the
Society:
“Total receipts, including balance for 1907..............$818.40
“Expenditures, OCLs oe. a ener OS
Tegal GS Fa Pasay eae ee 632.33
— 709 .96
Gashtonunamdeebec, NowlOORe. ts a Re ae eee aoe $108 . 44
(Signed) J. CHESTER BravDLey, For the Committee.
The following report from the auditing committee was
accepted and the committee was discharged :
‘The Committee appointed to audit the accounts of the Secretary-
Treasurer and the Managing Editor of the Annals respectfully report
that they have examined the same, compared them with the accom-
panying vouchers and found the same to be correct.”
(Signed) Puri Py CAryeRrn.
Dec. 11, 1908. F. M. WEBSTER.
Mr. Viereck moved to ask the Committee on Nomenclature
to define what constitutes a species and variety, taking into con-
sideration what the ornithologists have done, and that they
bring the same up before the Commission on Nomenclature of
the International Congress of Zoology one year before the next
meeting of that Congress.
Adjournment.
At eight P. M. in the assembly room of McCoy Hall, the
annual public address was given before the Society by Dr. E. B.
Poulton, Hope Professor of Zoology in the University of Oxford,
on ‘‘Mimicry in the Butterflies of North America.’’ The speaker
was introduced by Vice President Smith. A large and apprecia-
tive audience was in attendance. The address was illustrated
by many beautiful colored slides. It will be printed elsewhere
in these ANNALS.
J. CHESTER BRADLEY, Secretary—Treasurer.
6 Annals Entomological Society of America [Mole
RESOLUTIONS
On THE DEATH oF Dr. Wn. H. ASHMEAD.
William Harris Ashmead, Naturalist, Honorary Fellow of
this Society, died on Oct. 17th, 1908. We, his colleagues, would
hereby give expression to our sorrow in the loss of a leader and
a friend—one who gave himself unsparingly to the advancement
of the science we cherish, who made himself by his zeal and
industry one of the foremost students of the Hymenoptera in the
world, and who furthered our progress by his: numerous and
valuable papers and by the prompt and generous aid he lent to
every student who asked his expert knowledge and assistance.
We gratefully acknowledge our debt to him, and we desire to
place on record this testimonial of our esteem for him as a man,
our pride in his successful career, our high regard for his scientific
work, and our sincere sense of irreparable loss at his passing away.
j.2He Comstock:
J. G. NEEDHAM,
J.C. BRADLEY:
Commuttee.
WILLIAM H. ASHMEAD,
Plate I,
DR. JAMES FLETCHER.
Plate Il.
1909] Proceedings 7
RESOLUTIONS
On THE DeatH oF Dr. JAMES FLETCHER.
WHEREAS, By the untimely removal of Dr. James Fletcher, the
Entomological Society of America has lost—the first by death—
one of its original Fellows, and former Vice President, who pre-
sided at the Chicago meeting one year ago; and
WHEREAS, Dr. Fletcher, by reason of his nobility of character,
kindness of heart, and zeal, tempered by good judgment, was
known and beloved among scientific men not only throughout
the whole of the United States and the Dominion of Canada,
but also abroad, as a careful, conscientious, scientific worker,
a true Christian and a thorough gentleman; and
WHEREAS, his death, almost in the prime of life, is a serious
loss to the applied Science of Entomology as well as of Botany
in America; therefore, be it
Resolved, That by and through these resolutions the members
of this Society express their grief over this loss to two nations,
of this truly fine, good man and colleague; and be it further
Resolved, That a page in the Annals of this Society be set aside
for the purpose of placing these Resolutions on record and that
the Secretary be instructed to send a copy thereof to the bereaved
family.
F. M. WEBSTER,
F. H. CHITTENDEN,
C2 Lb. MARVATT,
Commiuttee.
8 Annals Entomological Society of America [Void
REPORT OF THE COMMITTEE ON NOMENCLATURE.
Your Committee desires to report that since its appointment,
four matters have been presented for its consideration. Of these,
one was the consideration of a particular case and was soon
settled. The second 1s still under consideration; the third it has
not as yet been able to take up, and the conclusions which the
Committee have reached upon the fourth case are herewith
presented.
The nomenclature of gall insects was referred to the Commit-
tee as the result of a paper by Dr. E. P. Felt presented at the last
meeting. The Committee is not unanimous on all points, but
considers it desirable to present the following:
Report on the Nomenclature of Gall Insects.
In the literature relating to galls and gall insects, there are
found several different kinds of description, accompanied by
names.
(1) Those relating to the galls only, with names intended to
apply to the galls, not to their inhabitants.
(2) Those relating to the galls only, but with specific names
referred to particular insect genera, and intended to apply to the
gall insects themselves, these being known at the time only from
their work.
(3) Those relating to the galls and the contained larvae, with
names proposed to be applied to the insects.
(4) Those relating to the galls and the adult insects bred
therefrom, and sometimes also to the larvae, with names proposed
for the insects in the usual manner.
(5) Those relating to the adult insects, the galls being un-
known, with names as usual.
I. It is agreed that in cases falling under No. 1, the names
proposed do not enter Zoological Nomenclature. It is also obvious
that in cases 4 and 5, the names are correctly proposed, and
available for use if otherwise in accordance with the International
Rules.
II. It is the opinion of the Committee that specific names
based on larvae (case 3) are available, and may be used.
III. With regard to the description of the gall, it is recog-
nized that it forms a valuable part of the diagnosis of any gall
insect, and that without it the recognition of the species may be
difficult or practically impossible, especially when the description
is not very detailed or precise. The Committee is willing to
accept a name based on the description of an adult or larva plus
It, PrarE Te
OL.
V
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1909} Habits of Wall Bee )
gall, even though the name would not be recognizable or of cer-
tain application were the account of the gall excluded from con-
sideration.
IV. With regard to names applied as in case 2, intended to
refer to the then unknown makers of known galls, it is the sense
of the Committee that whenever possible, these names should be
adopted.
V. The Committee is not wholly in agreement as to whether
it is obligatory to maintain names (if otherwise valid) proposed
as in case 2; or whether, when they are maintained, the original
author and date should be cited, or the author and date of the
publication in which the insect itself is first described. The
majority of the Committee, however, is against the obligatory
recognition of names accompanied by descriptions of galls only,
and holds that when these are adopted, they properly enter no-
menclature at the time of the description of the insect itself.
VI. The Committee agrees, that whatever may be the ulti-
mate ruling on the last point, there are many practical difficulties
in the way of recognizing names proposed as in case 2, so that
even were such names held to be available, many of them would
have to be rejected as of uncertain application. It is perfectly
clear that no rules will absolve an author from using his critical
judgment in the several cases that come before him; and after
the rules have declared a name available from their standpoint, it
may be a long way to availability from the standpoint of practical
identification.
The Committee is greatly indebted.to Dr. C. W. Stiles, the
Secretary of the International Commission on Zoological Nomen-
clature, for a full and luminous discussion of the matters in dispute.
H.'T. FERNALD,
LRDMAs COCKERPLL,
BAP PP arr:
A NOTE ON THE HABITS OF THE WALL-BEE,
CHALICODOMA MURARIA.
J. H. Comstock.
On the banks of the Nile, about 4oo miles above Cairo,
there stood for many centuries the city of Dendera, one of the
most ancient and most famous cities of Egypt. According to a
legend a temple was built here in the time of the Ancient Empire,
7. e., more than two thousand years before Christ. Of this temple
we know little; but about 2000 years ago the Egyptians built
IO Annals Entomological Society of America [Vol. IT,
another temple here, which is one of the better preserved of the
Egyptian temples.
The old city of Dendera has passed away. The houses built
of brick of Nile mud have crumbled to dust, and the sands of the
desert drift over much of the formerly cultivated fields. But
fortunately the drifting sands buried and preserved Dendera’s
magnificent temple.
Modern archeologists have excavated this temple. The
magnitude of the labor involved in this undertaking can be seen
by a glance at Figure 1 in Plate III, which gives a general view
of the temple and of the mounds of earth that have been removed
from it. The object of this note is to.show that, to a great
extent, the prodigious work of excavation that has been accom-
plished is being undone by an insect.
It seems evident to a layman, I know little of Egyptology,
that in the study of these ancient temples there are two main lines
of investigation: first, the study of the general architectural fea-
tures of the temples; and second, the reading of the inscriptions
with which the walls and the columns of the temples are covered.
Of these the second is certainly not less important than the first.
The temple of Dendera has been largely exposed, so that
its architectural features can be studied; but the inscriptions on
its walls are being rapidly buried beneath a layer of the cement-
like nests of the wall-bee, Chalicodoma muraria, as is shown in
Figure 2 of the Plate.
If future students and travelers are to see these inscriptions,
it will be necessary to carefully remove this layer of nests; and
as the nests are quite firm in texture, the labor involved will be
considerable.
Entomologists are not accustomed to look upon this bee as
a noxious insect. The studies of instinct and the embryological
investigations that have been pursued upon it in Europe have
made it a favorite subject of study; and I have not read of its
doing serious injury there. But at Dendera it is certainly a pest;
and we are warranted in suggesting means of destroying it.
The most practicable method of doing this that occurs to
me is the destruction of all of the nests at frequent intervals so °
that no bees can develop. The duration of the larval state in this
locality at the different seasons of the year should be determined
as a basis for determining the frequency of the removal of the
nests.
CONTRIBUTIONS TO OUR KNOWLEDGE OF THE ANATOMY
AND RELATIONSHIPS OF SPIDERS.
ALEXANDER PETRUNKEVITCH, Pu. D.
INTRODUCTORY REMARKS. TERMINOLOGY.
Walckenaer may have been right when in 1837 he pointed
with pride to the long list of works on spiders already in existence
and said that if Pallas could only see it, he would surely no
longer reproach naturalists for neglecting this interesting group
of animals. Unfortunately the reproach of Pallas may be
repeated now more justly than ever before. It is true that a
quantity of excellent work has been published since the time of
Walckenaer, much more and much better work than before. But
the group remains a neglected one especially if compared with
such favorites as insects or vertebrates. When two years ago
I began to gather material for a manual on spiders I soon became
aware that our knowledge of their anatomy and embryology is
entirely inadequate. Not only the nervous system and the
sense organs, but even the external sexual organs from which
the characters for the identification of species are chiefly derived
need thorough re-examination. This sad discovery stimulated
me to new research and the present series of articles is a result of
it. I shall omit all reference to literature which will be exten-
sively treated in the manual I hope to be able to complete and
to publish some day. Only such papers will be mentioned as are
absolutely necessary to the understanding of the subject. More-
over to avoid loss of time and to make the reading easier I shall
describe things as I found them during my investigations, with-
out making constant mention as to what is new and what was
already known. Those who know the literature will easily sep-
arate the new from the old, while those who are not so well
acquainted with it will be glad to have the subject treated in a
short and comprehensive manner. I need scarcely add that
everything described by me in the present series of articles is
based on careful and extensive study.
Concerning the methods I have applied during my investiga-
tions I have little to say. They are the ones commonly used by
zoologists and anatomists, i. e., dissecting with the aid of scissors,
scalpels and needles for macroscopic examination, at times aided
by in toto staining and injections; mounting of whole spiders or
ee
12 Annals Entomological Society of America [Volk Tt:
parts in canada balsam, glycerin or some other medium; and
finally sectioning with the microtome after applying the usual
reagents. Used alone none of those methods is quite reliable.
But when one is controlled by the others and when the number
of specimens and forms studied is sufficiently great, one gets a
clear idea of relations which would otherwise entirely eae obser-
vation.
To one point I desire to call particular attention. The ter-
minology in this group is in as bad a shape as in any other group
of animals. Even the names for the three planes of the body
have not found universal acceptance in their application to the
different groups of the animal kindgom and what is still worse the
same term has often more than one entirely distinct application.
Thus the term TRANSVERSE plane is used by different authors to
designate each of two separate intersecting planes, which results
in terrible confusion. In the case of appendages the terms upper
and lower, anterior and posterior, internal and external have
also been applied to the same structures, making it at times
extremely difficult to understand the author’s exact meaning.
To avoid all misunderstanding I shall therefore give in this intro-
duction the terms as I will use them throughout the entire series
of articles as well as in my papers on systematics, the first of
which will be ready for print in the near future. Whenever pos-
sible I preferred new terms to old ones, not out of desire to aug-
ment the number of terms of which there are already more than
needed, but to enable the reader to know at once and exactly
their meaning. Some of these terms are cumbersome and pos-
sibly could be replaced by better ones. However, clear and
exact terms are in my opinion to be preferred to those which
although easily remembered are at the same time easily confused.
The three determining planes of the body, applying to all
animals I will call as follows
1. THE PLANE OF SYMMETRY (called usually the sagittal or
chief plane). It divides the body into two symmetrical parts,
and not only in arthropods but even in echinoderms and coelen-
terates there is only one plane of symmetry. All planes that
run parallel to it I will call parasymmetrical.
2. THE SYNAXONIAL PLANE (called often the frontal, and By
Claus and others wrongly the transverse plane) is a plane inter-
secting the plane of symmetry at right angles in such a manner
that their line of intersection forms the chief axis of the body
1909] Anatomy and Relationship of Spiders 13
or the axis of complex symmetry, if I may use a term accepted in
crystallography. The position of the synaxonial plane in coelen-
terates and echinoderms is quite exact since the chief axis itself
is determined by the points of intersection of the radii. In
Arthropods in which the axis is not exactly determined by the
structure of the body itself, we may accept as the synaxonial
plane, the plane which divides the tergites from the sternites.
In the case of spiders it is the plane which separates the cepha-
lothorax from the sternum. Any plane parallel to this plane
I will call parasynaxomial.
3. THe DiaxoniaL PLANE (usually called the transverse
plane) is a plane intersecting the former two (and the chief axis)
at right. angles. Any plane which does this is a diaxonial or
transverse plane.
An exact terminology may be derived from these terms for the
different planes, surfaces and edges of appendages, such as legs,
antennae, palpi, chelae, etc., since all these are built on the prin-
ciple of bilateral symmetry. In consequence they have all three
determining planes of their own. Such terminology is especially
important in systematics. "To comprehend the new terms clearly,
one must imagine all legs as growing out of the sides of the ani-
mal at a right angle to its chief axis and the chelae and mouth-
parts as parallel to it. (Fig. I.) For the sake of convenience
we may drop the prefix ‘‘para,’’ since this does not impair clear-
ness. As will be easily understood from an examination of the
diagram, the terms apply to both sides of the body equally well,
so that it is not necessary to mention ‘‘left”’ or ‘‘right”’ and yet
no confusion can result. The same term applies to a surface
whether it be a leg of a mammal or of a spider, whether a front
or a hind leg.
1. THE EPISYNAXAIL SURFACE (should be epiparasynaxonial)
commonly called ‘‘dorsal.’’ The term dorsal is however object-
ionable since the articulation of the limbs is frequently such that
the episynaxial surface is not really dorsal in position, as for
example in the crabspiders.
2. THE HyposyNAXIAL SURFACE (should be hypoparasynax-
onial) commonly called ‘‘ventral,’”’ the latter name being object-
ionable on the same ground as ‘“‘dorsal.’’ The terms dorsal and
ventral I will apply to the body only, not to the appendages.
3. THE PROSYMMETRICAL SURFACE (should be proparasym-
metrical) is the lateral surface nearest to the anterior end of the
14 Annals Entomological Society of America [Wol--it,
chief axis of the animal (Fig. I. pro). It corresponds therefore
to the “‘inner”’ surface of the front legs, or to the ‘‘outer”’ surface
of the hind legs in spiders. It also corresponds to the ‘‘inner’’
surface of the chelae in Theraphosidae called “‘anterior”’ in true
spiders.
4. THE RETROSYMMETRICAL SURFACE (should be retropara-
symmetrical) is the lateral surface nearest to the posterior end
of the chief axis of the animal. It corresponds to the “‘outer”’
side of the front legs or to the “‘inner’”’ side of the hindlegs.
(Fig. I. retro); In the chelae it corresponds to fhe, souter™
surface Theraphosidae, or ‘‘posterior”’ in true spiders.
The four edges or lines of intersection of the four surfaces
mentioned in those cases when the appendages are not cylindri-
cal in form may be called “‘epipro”’ (Fig. 2. ep), ‘‘epiretro”’ (Fig.
2, er), ‘‘hypopro’’ (Fig. 2, hp) and “‘hyporetro”’ edge, and the
middle lines or the lines of intersection of the three determining
planes with the four surfaces mentioned may be called the upper
and lower, pro and retro middle line. In the case a structure or
edge is oblique as the margins on the chelae for example, it may
be simply designated by the word pro or retro. Thus the ‘‘marge
inferieur’’ of Simon’s terminology, called by Banks ‘‘the row
behind the fang” and by Montgomery ‘‘the posterior margin”’ is
according to my terminology the Retromargin; while the “‘marge
superieur’’ of Simon or the anterior margin of Montgomery is the
Promargin. There can be no confusion if we apply this terminol-
ogy consecutively, no matter what the actual position of the limb,
if we only remember that each limb has only one plane of sym-
metry and that the entire terminology is based on the correla-
tion of planes and their homology in the left and right side of
the animal.
I. The Structure and the arrangement of har on the legs
of Pholcus phalangoides Fuss.
Little attention has been paid to the hair covering the legs
of spiders. This is deplorable since many a valuable character
may still be derived from its study, as the investigations of Dahl
on the trichobothria have proven. They are of great help in the
separation of families although their value should of course not
be overestimated. I do not know whether the hair will even-
tually furnish characters of value for the separation into genera,
but it has been already applied for the separation into species of
1909] Anatomy and Relationship of Spiders 15
the genus Latrodectus and it certainly will help to establish
relationships between forms which would otherwise remain sep-
arated in the system.
While studying the muscular system and articulations in
Pholcus I was struck by the regularity of the rows of hair on the
legs. A comparison of many specimens showed that this is a
constant character. The leg has not the form of a perfect cylin-
der, but shows in a cross-section a circle with flattened sides, repre-
sented diagrammatically in Fig. 2. There are in all 8 rows of
hair; two rows on the episynaxial surface (esp and esr), one on
the middle pro (mp) and one on the middle retro line (mr),
and four rows on the four edges (ep, er, hp, hr). We find the
same arrangement on each segment of all legs beginning with the
femur; but the tarsus of the fourth leg in both sexes has a ninth
row of hair on the middle hypo line. The onychium of each tar-
sus is surrounded in all legs by 8 bristles, two of which are unpaired
and belong to the tarsus. The structure of these bristles may
best be understood from the drawing. (Fig. 4.)
The hair forming the eight rows on the femur and tibia is of
uniform structure. It is represented in Fig. 5, in its entire length
and a part of it on a larger scale in Fig. 6. It is characterised
by the spiral row of small spines reaching a little beyond its
middle. Each hair is inclined towards the distal end of the limb
at an angle of from 27-30 degrees. Ina female of 2 mm. cepha-
lothorax length the hair on the femur marked in Fig. 2, with the
letters esp and esr is about 0.8 mm. long, while that forming the
other four rows is only 0.57 mm. long. In the tibia all rows con-
sist of hair of the average length of 1.0 mm. There are no tric-
chobothria on the femur and tibia, but on the latter are some
six or seven extremely small hairs without any special structure,
measuring about o.o6 mm. The hair of the metatarsus is con-
siderably shorter than that of the tibia, but still of the same
structure in its proximal two thirds. The longest hair is at the
base of the metatarsus measuring in the above individual o.4 mm.
Towards the distal end of the metatarsus the hair becomes grad-
ually shorter and changes structure. It measures now only about
0.28 mm. loses the spiral of small spines, while the two basal
spines—the only ones remaining—become considerably stronger.
(Fig. 7.) There are besides some six or seven scattered hairs
measuring about 0.12 mm., simple at the proximal end of the
segment and toothed at its distal end, where it looks as represented
16 Annals Entomological Society of America [Volitl,
in Fig. 11. This hair may be bent anteriorly or erected more than
the common hair. Its teeth are directed away from the tip of
the limb. Similar hairs, only shorter and with longer teeth are
found on all tarsi,about one dozen in number (Fig.1o). The hair
forming the eight rows on the tarsi is of the same structure as
that at the distal end of the metatarsus, but also smaller, measur-
ing from o.2-0.28 mm. and with stronger basal spines (Fig. 8)
to which two more are sometimes added. The fourth leg in both
sexes has besides, as already mentioned, a ninth row of specially
structured hairs. They occupy the hypomiddle line and are
from thirty to thirty-four in number. They have hooks on one
side and little spines on the other (Fig. 9) and are so arranged
that the spines are directed distally towards the tip of the limb.
Although each hair measures only 0.12 mm. this row doubtless is
homologous with the comb of the Theridiidae, since the position of
the row is in each case the same and the structure of the hair, if
we disregard its length, very much alike. This brings the family
of the Pholcidae, originally regarded as a subfamily of the Theri-
didae and later separated from the latter family, into closer rela-
tion with it again. Such opinion is further substantiated by the
fact that other representatives of the Family Pholcidae show simi-
lar conditions. In Spermophora meridionalis the row consists of
16 hairs one of which is represented in Fig. 12. In two species of
Modisimus from Jamaica I find this row represented by hairs of
the type found in the other eight rows, but the distal four hairs
show distinct teeth or spines characteristic for the combhair.
In Pholcus tipuloides the whole row differs from that of Pholcus
phalangoides, resembling more the lower end bristle of this latter
species and thus constituting a character in itself sufficient to keep
the two species apart. On the other hand neither Hypselistes
(Erigone) florens, nor any of the five common species of Linyphia
examined, show anything of the kind.
II. On the Muscular System of the legs. .
The muscular system of the legs in spiders has been studied
several times. The last paper of importance was that by Paul
Gaubert in the Annales des Sciences Naturelles, Vol. 13, 1892.
Yet neither his studies, nor those of Borner have brought the
question to a conclusion. Moreover they contain errors. Thus
one of the muscles of the patella was entirely overlooked and
another in the tibia wrongly described and in consequence its
function misunderstood.
1909] Anatomy and Relationship of Spiders 17
I used as material transparent legs of small spiders. They
can be studied in life, or after enclosure in glycerin, or by fixing
the whole leg in picric acid which stains intensely and perma-
nently yellow the muscles and by enclosing it then in canada
balsam. Side views were controlled by dorsal and ventral views
of corresponding segments. In some cases sections were neces-
sary and the paraffin-method was used. Besides small spiders I
have also studied the muscular system in several very large spi-
ders, such as Heteropoda venatoria, Ctenus malvernensis sp. n.,
Lycosa carolinensis, Pachylomerus nidulans and unidentified
species of Eurypelma. Inallthese the muscles can be isolated with
the aid of dissecting instruments even without a magnifying glass.
Coxa. Enclosed in the coxa are five muscles one of which
really belongs to the trochanter and will be described with the
other muscle of that segment. The four muscles belonging to the
coxa are: 1. WM. flexor trochanteris (Fig. 3, fl. tr.) This muscle
arises from the episynaxial surface of the coxa and partly from
the chitinous septum which occupies the synaxial plane of the
coxa and divides its anterior two thirds into an upper and lower
half. The muscle is inserted with a tendon into the proximal
epi-edge of the trochanter. 2. M. extensor trochanteris (Fig. 3.
ex. tr.), the antagonist of the preceding arises from the hypo-
synaxial surface of the Coxa and partly from the septum. Its
tendon is inserted into the proximal hypo-edge of the trochanter.
3. M. promotor trochanteris, a broad muscle with parallel fibres
which arise from the prosymmetrical surface of the Coxa and are
inserted without tendons into the proximal prosymmetrical edge
of the trochanter. 4. M. retractor trochanteris (Fig. 3. r. tr.) the
antagonist of the promotor occupying the symmetrical position
on the retro-side.
TROCHANTER. ‘Two muscles are enclosed in the trochanter.
1. VM. flexor femorts longus (Fig. 3. fl. f. 1.) a weak muscle which
arises from the coxal septum and is inserted by means of a tendon
into the proximal epi-edge of the femur. Gaubert, who calls this
muscle “‘abaisseur,’’ describes it as arising from the inferior
(Hyposynaxial of my terminology) surface of the coxa, not from
its septum. I think that in some cases I have seen fibres running
in that direction, but I am not positive about it. The function
of this muscle is certainly not that of the common flexor but it is
not an ‘‘abaisseur’’ (depressor), since it probably helps the flexor
bilobatus to lift the femur and at the same time tends to bring
18 Annals Entomological Society of America [Melt
the whole femur nearer towards the trochanter. 2. M. flexor
femoris bilobatus (Fig. 3. fl. f. b.) consists of two symmetrical
halves arising from the hyposynaxial surface of the trochanter.
Its numerous short tendons are inserted close together into the
same edge with the flexor longus.
FemMuR. Enclosed in the femur we find two muscles closely
resembling those of the trochanter. 1. WM. flexor patellae longus
(Fig. 3. fl. p. 1.) has two proximal tendons one of which arises
from the distal end of the hyposynaxial surface of the trochanter
and the other from the corresponding proximal end of the femur.
The fibres soon form one spindle-like muscle which is inserted
with a long tendon into the proximal hypo-edge of the patella.
2. M. flexor patellae bilobatus (Fig. 3. fl. p. b.) is in every respect
homologous with the corresponding muscle of the trochanter, but
as the flexor femoris raises the femur, while the flexor patellae
lowers the patella together with the tibia, the two homologous
muscles arise from the opposite surfaces. It is the articulation
of the segments that determines the position of muscles. The
musculus flexor patellae bilobatus consisting of two symmetrical
halves arises therefore from the entire episynaxial surface of the
femur and its numerous short tendons are inserted close together
into the proximal hypo-edge of the patella on both sides of the
long flexor.
PaTELLA. The articulation of the patella with the tibia is
such as to allow of scarcely any motion. Notwithstanding this
we find three muscles in the patella. Two of these are symmetri-
cal and consist of parallel fibres arising from the episynaxial and
the pro- and retrosymmetrical surfaces of the segment. Each
separate fibre is inserted apparently without a tendon into the
proximal pro- and retro-edges of the tibia. These two muscles
are: 1, M. promotor tibiae, and 2, M. retractor tibiae (Fig. 3. r. t.)
The third muscle which is also to all appearances functionless 1s
3, M. flexor tibtae. It arises from the proximal end of the episyn-
axial surface of the patella and is conicalin shape. Its long cylin-
drical tendon is inserted into the middle of the proximal hypo-
edge of the tibia. This muscle is entirely overlooked by previous
investigators although it may easily be found in spiders of differ-
ent families and all sizes.
Tipra. The muscles of the tibia are in every respect homolo-
gous with those of the femur. A number of fibres arise from the
proximal surface of the patella without however forming a tendon,
Tg09| Anatomy and Relationship of Spiders Ig
and join in the tibia those fibres which arise by means of a tendon
from the proximal hypo-edge of this latter segment. Together
they form the 1.—WM. flexor metatars: longus (Fig. 3. fl. m. 1.)
which is inserted with a long tendon into the middle of the prox1-
mal hypo-edge of the metatarsus. 2, W/. flexor metatarsi bilobatus
(Fig. 3. fl. m. b.) is composed of two symmetrical halves and arises
from the entire episynaxial surface of the tibia. Its numerous
short tendons are inserted closely together on both sides of the
tendon of the long flexor. The description of this muscle given
by Gaubert is entirely wrong and I am not able to comprehend
what has brought him to the conclusion that its point of
insertion is on the dorsal (episynaxial) surface of the metatarsus
and that this muscle is therefore an extensor. There is no exten-
sor of the metatarsus.
Meratarsus. Enclosed in the metatarsus are two muscles,
one of which is the antagonist of the other. 1, WM. flexor unguium
(Fig. 3. fl. u.) arises from the proximal surface of the metatarsus
and receives additional fibres which arise from the distal end of
the episynaxial surface of the tibia. Its long, thin tendon tra-
verses the metatarsus and tarsus and is inserted into the base of
the claws. 2, VM. extensor unguium (Fig. 3. ex. u.) arises from the
proximal epi-edge of the metatarsus and is inserted by means of
a long and thin tendon which runs above and parallel to that
of the flexor into the base of the claws above the latter.
Tarsus. The tarsus has no muscles and encloses merely the
two tendons above mentioned.
In reviewing the muscular system of the legs in spiders we
observe that its characteristic feature is the absence of extensors
in the majority of segments. Only the coxa and the metatarsus
enclose extensors. The function of the extensors is evidently
transferred in the other segments upon the hypodermis forming a
thin elastic membrane over each joint. The homology of the
muscles enclosed in the femur and tibia is too apparent to admit
of doubt, and I believe that it can be extended to the muscles
1. In the second volume of his text-book of experimental zoology, which has
just appeared, Przibram reproduces Frédéricg’s diagram of the muscular system
of a spider-leg. Evidently misled by Garber’s old interpretation of the muscular
system of appendages in Arthropods, he figures a flexor and an extensor in every
segment of the limb. However, even a superficial examination of the articulations
cannot fail to show that his supposed extensors are in reality flexors. If he had
noticed that the axes of the patellar and metatarsal articulations lie not on the
ventral side of the limb, but in its episynaxial (dorsal) surface, he would have
escaped this error.
20 Annals Entomological Society of America [Vol JLr:
enclosed in the trochanter. Such conception is certainly true in
the case of the flexor bilobatus. As for the flexor femoris longus,
some doubt may be expressed as to its homology with the corres-
ponding flexors of the patella and metatarsus. But the short-
ness of the trochanter and the difference in articulation may be
held responsible for the fact that the flexor femoris longus arises
from another surface than its homologs. The muscles of the pa-
tella are doubtless homologous with those of the coxa with that
exception that there is no extensor tibiae. We must however
remember that an extensor tibiae would be of still less use than
the flexor, since the patella has a special tooth at its distal end
interlocking with the tibia. This interlocking device prevents
the tibia from being raised apart from the patella, the only func-
tioning “‘knee”’ articulation being that between the femur and
the patella.
EXPLANATION OF THE FIGURES (PLATE IV.)
1. Diagram of a spider showing the axis of complex symmetry, AP, and
the planes of symmetry of the legs, pd, and of the chelae. The second and
third legs on the right side are removed. ‘The figure represents the dorsal sur-
face of the spider and the legs are extended in such a manner that the surface
of the paper represents the episynaxial surface.
2. Diagram representing a cross section of the leg of Pholcus phalangoides.
EH, the plane of symmetry. It is understood that the observer sees the cross
section in looking from the body of the spider towards the end of the leg. In
consequence it is evident from the letters accompanying the diagram that it is
across section of arightleg. Ina left leg the two halves are reversed. ESP, pro-
episynaxial; ESR, retoepisynaxial; EP, apipro, ER, epiretra; MP, middle pro;
MR, middle retro; HP, hypo pro; HR, hyporetro hair.
3. The muscular system of a spider leg. fl. tr., flexor trochanteris; ex. tr.,
extensor trochanteris; r. tr., retractor trochanteris. Its antagonist, the M. pro-
motor trochanteris, runs parallel to the retractor and cannot therefore be repre-
sented on the diagram. One must imagine that it was removed for the purpose
of showing the other muscles. fl. f. 1., flexor femoris longus; fl. f. b., flexor fe-
moris bilobatus; fl. p. 1., flexor patellae longus; fl. p. b., flexor patellae bilobatus;
fl. t., flexor tibiae; r. t., retractor tibiae. Its antagonist the M. promotor tibiae
runs parallel to the retractor and could not be represented in the diagram.
fl. m. 1., flexor metatarsi longus; fl. m. b., flexor metatarsi bilobatus; flu., flexor
unguium; ex. u., extensor unguium- tn. f., the tendon of the flexor; tn. e., the ten-
don of the extensor.
4. The bristles surrounding the claws of Pholcus phalangoides. The
heavy upper and lower bristles are single. Three bristles corresponding with
the three other bristles of the figure are situated on the other side of the claws
and can not be represented.
5—11. Hairs on the legs of Pholcus phalangoides. 5, on femur, patella,
tibia and the two proximal thirds of metatarsus. 6, a part of the same hair
stronger magnified. 7, a hair from the distal end of the metatrasus; 8, a hair
from the tarsus. 9, the combhair of the fourth tarsus. 10, one of the few
erect, scattered hairs on the tarsus. 11, the same from the metatarsus. 12, one
of the combhairs of Spermophora meridionalis.
ANNALS E. §. A. VoL. II, PLATE IV. -
aD eh FigT Fig 8. Fig. 9. Fig. 10. Fig A. Fig. IL.
Alexander Petrunkevitch.
A MONOGRAPHIC CATALOGUE OF THE MYMARID GENUS
CAMPTOPTERA FOERSTER, WITH DESCRIPTION OF
ONE NEW NORTH AMERICAN FORM.
A. ARSENE GIRAULT, UNIVERSITY OF ILLINOIS.
In recording the discovery of another new form of this genus
from North America, I have thought it helpful to bring together
the original descriptions of the few known species, and to sum
up our knowledge concerning the group. What we know of the
genus, however, amounts practically to very little. But three
species have hitherto been described, none of them any too well.
HISTORY OF THE GENUS.
The genus Camptoptera was originally characterized by Arnold
Foerster in 1856 (Foerster, 1856, p. 116) as follows:
“28. Mymaroidae. Die mir bekannten Gattungen lassen sich
nicht alle, ohne die G@ und 2 zu trennen, in eine einfache und klare
Uebersicht bringen. Das nachfolgende Schema wird daher einige
Gattungen doppelt auffthren:
a. Die Tarsen finfgliedrig.
b. Der Hinterleib deutlich gestielt.
c. Die Fuhler beim % zehn-, beim 9 neungliedrig.
Camptoptera 7.*)
cc. Die Fihler beim @ dreizehn-, beim 9 eilfgliedrig.
Ooctonus Hal.
bb. Der Hinterleib sitzend oder fast sitzend.
a. Die Giseere: pasklGe
Further on (Id., p. 119), he gives the characters of the group
as follows:
‘“Unter dem Namen Camptoptera habe ich eine pentamere Gattung
aufgestellt, welche durch ihre winzige Form sich nur mikroskopisch
bestimmen lasst. Da ich dieses Thierchen erzogen habe und lebend
untersuchen konnte, so will ich mich hier etwas weitlaufiger darttber
verbreiten. Der Gattungscharakter lautet:
Tarsen ftinfgliedrig; Fuhler beim @ zehn-, beim @ neungliedrig;
Hinterleib kurz gestielt; Flugel schmal, mit einem breiteren Absatz
nahe an der Basis und hinter demselben gebogen.”’
A description of the species (papaveris) upon which the genus
was founded then follows in the next paragraph; although this
“*)Camptoptera von xaumrrds, 7, 6v, gekriimmt, gebogen und mrepéy, 76,
der Fligel. Die Vorderfltigel sind in dieser Gattung ziemlich stark gebogen.”’
22
1909 | The Mymarid Genus Camptoptera 23
LS)
2
species is not designated as the type, under the code it becomes
such (monotpyical genus, Stiles, tg05), and was formally desig-
nated as such by Ashmead (1904).
Still later on, in the same work, under the heading Nachtrag,
Foerster (1856, p. 144) proposed to change the name of the
genus from Camptoptera to Pteroclisis on the ground that ‘‘da
bereits durch Pres! der gleichbedeutende Name Camptopteris in
die fossile Flora eingefthrt wurde.’ This proposed change right-
ly was not accepted by later authors and under both the present
code (Stiles, 1905, p. 10, independence from botany; p. 25, page
precedence) and the law of priority, the second name is aban-
doned. The incident is stated as a matter of history.
The genus has received little or no attention since its estab-
lishment by Foerster, but later authors (Provancher, 1889; Cres-
son, 1887; Ashmead, 1904) have persisted in spelling it Camp-
totera.
The former author gives the following generic description
(Provancher, 1889):
““Gen. 11. Camptotére. Camptotera, Forst.
Antennes insérées sur le front, non presde la bouche. Abdomen
plus ou moins arrondi sur les cétés. Ailes posté-térieures trés
petites, presque linéaires, sans nervure médiane. Abdomen
petiole; tarses de § articles:
Une seule espéce rencontrée.”’
DISTRIBUTION OF THE GENUS.
This genus is now represented in Europe and North America;
papaveris, the European species was described from Germany;
Provancher described clavata from the vicinity of Quebec, Canada,
while metotarsa Girault was described from Virginia and pulla
Girault is now described from Hlinois.
HOST RELATIONS OF THE GENUS.
Of the species clavata, metotarsa and pulla nothing is known
about their hosts, all of them having been described from captured
specimens; however, papavarts, says Foerster, “‘erzog ich vom 12.
bis 18. August 1851 aus Mohnkapseln von Papaver Rhoeas und
dubium, worin die Gallen vou Aulax Rhoeadis sich befanden.”’
Foerster (1856, p. 120) further adds in the next paragraph: ‘‘Ob
nun die Larven der Gattung Camptoptera an den Larven oder in
den Eiern der Gallerzeuger, hier des Aulax Rhoeadis Hart. para-
sitisch leben oder die Cecidomyia angreifen, oder gar die wbrigen
24 Annals Entomological Society of America Vols i:
Parasiten vernichten, daruber habe ich mir keinen Aufschluss
verschaffen kénnen, wird aber jederfalls von Interesse sein und
hoffentlich auch von anderer Seite nicht ohne Beachtung bleiben.”
The species was reared from poppy capsules containing a cynipid
gall and from which were reared also a cecidomyiid and three other
species of Hymenoptera; its host relations are therefore problem-
atical and it would be hazardous to attempt to connect it with
any of the possible hosts mentioned. This being the case, it is
seen that at present, of the four species of the genus, nothing is
known concerning their economy in nature. De Dalla Torre
(1898, footnote) on the contrary makes Foerster’s rearing a defi-
nite one, giving the host as Cynips papaveris (Perris) Mayr, and
the authority as Rondani. This Cynips is synonymic with
Aulax rhoeadis. The assumption in regard-to this host is all the
more arbitrary because of what Foerster himself states in regard
to the rearing. Kaltenbach (1874) also gives Foerster’s rearings.
Family Mymaridae.
Subfamily Gonatocerinae.
Tribe Ooctonini.
Genus Camptoptera Foerster, 1856, pp. 116, 119, 144.
(Type:—Camptoptera papaveris Foerster).
1. Camptoptera papaveris Foerster.
Foerster, 1856, pp. 119-120. Kirchner, 1867. Snellen van Vollenhoven,
1873. De Dalla Torre, 1898, p. 431. Ashmead, 1904, pp. 362, 369.
‘*Braunlich, der Schaft mit dem Stielchen, der Hinterleibsstiel und
die Beine gelb; die Fusse fast unmerklich dunkler als die tbrigen
Theile der Beine. Das & hat die Geisselglieder alle ungefahr gleich
lang, jedoch so, dass die vier bis flinf ersten Glieder fast unmerklich
langer erscheinen, als die folgenden; das letzte ist mit einer sehr feinen
pfriemenformigen Spitze versehen. Die Futhler stehen hart am
innern Augenrande. Beim 9 ist das erste Geisselglied etwas langer
als das Stielchen, das zweite viel langer als das erste, das dritte etwas
kurzer als das erste und das vierte sogar kurzer als das dritte. Die
beiden folgenden unter sich genau von gleicher Lange, einzeln
genommen wenig, aber doch merklich kttrzer als das dritte. Das
letzte Glied bildet eine ziemlich stark verlangerte, fast elliptische
Keule von der Lange der drei vorangehenden Glieder. Die beiden der
Keule vorhergehenden Glieder sind auch ein wenig dicker als die
ubrigen Geisselglieder.
Camptoptera Papaveris m. @ 9, die einzige Art dieser Gattung,
erzog ich vom 12. bis 18. August 1851 aus Mohnkapseln von Papaver
Rhoeas und dubium, worin die Gallen von Aulax Rhoeadis sich
befanden. Aus denselben Kapseln und zu derselben Zeit erzog ich
ferner: Pteromalus Papaveris m. @ 9. Cecidomyia Papaveris
to
Un
1909] The Mymarid Genus Camptoptera
=>
Winn.**)* Pezomachus Papaverism. 3 92 (das @ ungefligelt!!!) Loch-
ites Papaveris m. o @ (wortber bereits oben Seite 44 berichtet
wurde!) und ein kleiner Encyrtus.
Ob nun die Larven der Gattung Camptoptera an den Larven oder
in den Eiren der Gallerzeuger, hier des Aulax Rhoeadis Hart. parasi-
tisch leben oder die Cecidomyia angreifen, oder gar die tbrigen Para-
siten vernichten, daruber habe ich mir keinen Aufschluss verschaffen
konnen, wird aber jedenfalls von Interesse sein und hoffentlich auch
von anderer Seite nicht ohne Beachtung bleiben.”’” pp. 119-120.
This description is rather brief, lacks detail and it 1s more or
less doubtful if the species could be again recognized from it, so
that I have drawn up another from a female specimen nicely
mounted in balsam by Mr. Frederick Enock of London and deter-
mined by an English authority; as no other valid specimens of
the species exist, so far as | am aware, this description definitely
fixes the species; it will be seen readily that it agrees in general
with the original, though drawn up before that description was
consulted.
Redescription of Camptoptera papaverts Foerster, female.
Female.—Length, 0.91 mm.; minute; visible to naked eye. General color
yellowish brown; legs and basal five joints of antennae paler; ocelli red; eyes dark;
vertexal carina present, usual. Fore wings slightly infumated at basal one-third,
normal, curved at the apical fourth, with the usual dilatation along the caudal
margin near base, the usual dusky yellow margins and with one principal row of
discal cilia in the distal half of the wing (excluding the ones along the margin,
dorsad), joined by two others on each side, near the cephalic and caudal margins
respectively, of the wings, in the apical fourth; proximad of the apical fourth of
the wing, these two lines of cilia become lost in the margins; venation dusky yel-
lowish, inconspicuous. Hind wings hyaline, with two main rows of discal cilia,
one on each side near the margins, in the distal half of the wing; marginal cilia
of hind wings nearly as long as those of the fore wings, and with the usual color-
less spot or area near their bases, forming the clear whitish path a short distance
beyond the distal end of the wing, and which follows the outline of the wing
margin at that point. Petiole apparently acute at its center, lateral aspect
(as in pulla) and with a clear spot at its base, dorsal aspect; posterior tibiae
slender, longer than the posterior tarsi; the joints of the tarsi subequal, the basal
joints inclined to be slightly longer.
Antennae longer than the body; scape convexly curved, about as long as
the combined lengths of the pedicel and Ist funicle joint; pedicel conic-ovate, its
cephalic margin serrate, thicker than the scape and much more so than the slen-
der first three funicle joints, but nearly one-half shorter than the first funicle
joint; the latter slender, cylindrical, slightly narrowed in the middle, one-third
shorter than funicle 2 and the next to the longest joint of the funicle; funicle 2
the longest funicle joint, long and slender, one-third longer than 1, longer than
the scape and about equal to or slightly longer than the combined lengths of the
next two joints (funicles 3 and 4), subequal to the club in length, but one-half
narrower; funicle joints 1—3 of the same width, narrow and cylindrical; funicle 3
a little over half the length of funicle 2 and slightly enlarged at the apex, longer
than either of the following joints and also the pedicel; next three joints (funicles
4—6) gradually enlarging to club; funicle 4 cylindrical oval, distinctly wider than
3 and one-fourth shorter, subequal in length to 5; the latter slightly wider; funicle
a. The footnote to which the asterisks refer is omitted here, having no
relevancy.—A. A. G.
26 Annals Entomological Society of America [Vol. II,
6 one-fourth shorter than the two preceding, ovate, subequal in length to the
pedicel, the shortest funicle joint; club ovate, subequal to the combined lengths
of the three preceding joints (funicle 4-6), and from one-third to one-half wider
than funicle 6, and from 2 to 3 times the width of funicles 1-3, longer and wider
than the scape; more slender in proportion to the other joints than usual.
(Fig. 1. Flagellum of female, the setae omitted.)
(From 1 9, 2-3 inch objective Bausch and Lomb.)
Fig. 1. Flagellum of Camptoptera papaveris Foerster female greatly enlarged.
é 5 > 5
Redescribed from a specimen beautifully mounted in balsam
by Mr. Frederick Enock of London, and being one determined by
an English authority and in the collection of Dr. L. O. Howard.
Very similar, excepting in coloration and wing characters, to
pulla Girault. The antennae are very similar.
2. Camptoptera clavata Provancher.
Provancher, 1889, p. 404. De Dalla Torre, 1898, p. 431.
‘“‘Camptotere en-massue. Cam/ptotera clavata, n. sp.
?—Long. .og pce. Noire, polie, brillante, les antennes et les
pattes jaune-orange. Face inférieure; antennes insérées sur un
rebord du tront, de ro articles GY eto, articles 9° Curssesigins
ou moins renflées; tarses de 5 articles. Abdomen pédiculé, le
reste formant une massue en pointe a l’extrémité.—Ste Gertrude.”’
This brief, general description is inadequate for recognition
of the species, which on this account is probably lost to science
and will have to be rediscovered and renamed. The whereabouts
of the type is unknown to me.
3. Camptoptera metotarsa Girault.
Girault, 1905, p. 91.
“Female. Length, o.90 mm.
Shining black, the abdomen polished black. Legs, excepting
the coxae and apical joint of tarsi, basal three-fourths of the scape,
and the venation, brown.
Head about as wide as the thorax, the eyes subovate, coarse,
and very dark red; ocelli inconspicuous. Head and thorax
coarsely shagreened, the latter short and convex, slightly longer
than abdomen; parapsidal furrows absent. Abdomen depressed,
short, oval. Legs long, the first tarsal joint long, the coxae
rather large and coarsely shagreened, the posterior femur with a
convexity or swelling in the middle of its upper margin. Wings
iridescent, very slightly clouded, the fore wings narrow, curved,
the marginal fringes rather long. Body nearly hairless.
1909] The Mymarid Genus Camptoptera 27
Antennae elbowed, about as long as the body, g-jointed.
scape very long, subclavate, narrow basally, about as long, or
not quite so, as the next three joints combined, or about five times
the length of the pedicel. Pedicel short, very much smaller,
obconic. Joints 3 and 4 slender, cylindric, 4 longer, about twice
the length of the pedicel, both slenderer than the scape and pedi-
cel; 5, 6, and 7 equal, columnar, stouter than 3 and 4, but much
shorter, yet far longer than the pedicel. Joint 8 equal to 5, 6
and 7, but slightly stouter. Club cylindric, not as long as the
scape, but much stouter, about equal in length to 6, 7, and 8 com-
bined.
From a single female captured while jarring peach at Arlington
Virginia, June 2, 1905. A rather large and striking species.
Type.—No. 8941, United States National Museum.’ 1
tagmounted. Male, unknown.
This description also is not as detailed as it might be. The
type, being an unique, has not been accessible, but Mr. J. C. Craw-
ford, of the United States National Museum, Division of Insects,
at my request, kindly furnished the following additional and
important descriptive details: The petiole is acute or barbed at
the lateral margins, as in pulla; the vertexal carina is present;
and there are four rows of discal cilia on the fore wings—one near
the cephalic margin; two, not in stright lines, along the middle
portion of the wing, one on each side of a darkened line on the
wing, the line representing an obsolete vein; and the fourth near
the caudal wing margin. I am also informed that the type
specimen is broken, the head and thorax together parted from the
abdomen, but remounted on the same tag.
40
4. Camptoptera pulla species nova.
Female.—Length, 0.84 mm. Minute.
General color uniformly greyish black (near cologne earth); scape, pedicel,
Ist funicle joint (less so) and legs (including at least the intermediate coxae, the
others not visible) all pale clay yellow; remaining antennal joints concolorous
with body, the first funicle joint intermediate in color, dusky yellow; eyes and
ocelli dark red; usual vertexal carina present; thorax longer than abdomen, the
latter subnapiform, widest and acute at proximal third and narrowing gradually
to a point caudad, and truncately oblique to the petiole, cephalad; petiole short,
acutely produced or barbed laterad just caudad of its center and with a small
circular whitish dot in the middle of its dorsal aspect at base. Body apparently
impunctate. Legs normal, the tarsal joints subequal, the proximal tarsal joint
not much longer than any of the others, but subequal to them. Wings normal,
longer than body; fore wings hyaline, slightly fumated near bas2, and with a
slight but usual dilatation along the caudal margin at proximal fourth, curved
at the apical fourth, and with moderately long marginal fringes which have the
usual clear path around the edge of the wing at apex; discal cilia numerous, about
from 3 to 4 longitudinal rows; venation dusky, inconspicuous; apical margin of
28 Annals Entomological Society of America [Vol. IT,
fore wing more pronounced. Hind wing fumated throughout, its petiole yellow-
ish; curved at about its center, linear, and with at least two longitudinal rows
of discal cilia, very close together and central. Ovipositor very slightly exserted.
Antennae 9-jointed; scape curved, normal, longer than pedicel, the latter
ovate, with the apical end truncate and serrated, and but two-thirds the length
of the first funicle joint. Funicle joints 1— 3 cylindrical, linear, 1 one-third short-
er than 2 which is longest, subequal to the scape, and twice the length of funicle
joint 3, the three joints filiform; funicle joints 4-6 cylindrical oval, wider, sub-
equal in length but 4 is somewhat longer than the others, which, howev er, are
somewhat wider than that joint: funicle 4 subequal in length to joint 38, joints 5
and 6 each becoming slightly shorter; club long, undiv ided, cylindrical ovate,
longer than scape or funicle joint 2, and about equal the combined lengths of the
three apical funicle joints. Setae present, but not very numerous and minute.
Antennae longer than the body. (Fig. 2. Flagellum of female, the setae
omitted. The club is a little too long in the drawing.)
ee ee
Fig. 2. Flagellum of Camptoptera pulla Girault, female greatly enlarged.
(From two specimens, 2—3 inch objective. Bausch and Lomb.)
Male.—Unknown.
Described from two females mounted in balsam and collected
on a window in a woodshed, Urbana, Illinois, July 15 and 17, 1908
(J. Douglas Hood.) Several other specimens were observed July
17, 1908, on the same window suspended in a spider’s web, but
because of their minuteness were lost.
Type.—Accession No. 39116, Illinois State Laboratory of
Natural History, Urbana, Illinois, 1 2 in balsam.
This species resembles closely in structure of the antennal
joints, the wings, and that of the body, the type of the genus,
Foerster’s papaveris but differs markedly in coloration, (papa-
verts being almost uniformly colored), in the number of discal
cilia of the fore wings, in the fumation of the hind wings, in the
longer, slenderer club, and in its habitus.
TABLE OF THE SPECIES.
A. Species yellowish brown, color uniform.
Joint 2 of the funicle the longest funicle joint; scape about as long as
the combined lengths of the two “following joints; fore wings with but a
single<prominent tow Of discal cilia agen sete. t: os cieteesee eee papaveris
B. Species more or less black.
a. Shining black.
Antennae and legs “jatne—crange (1. 2 2 ee ee clavata
Antennae black, excepting basal three-fourths of the scape: legs
brown, excepting coxae; joint 4 of funicle the longest funicle joint;
scape about as long as the three following joints; fore wings with 4
longitudinal rows of discaliiciliameeenre seers ene metotarsa
b. Greyish black.
Antennae concolorous, excepting first three joints, which together
with the legs, are pale clay yellow. Joint 2 of the funicle the longest
funicle joint; scape not quite as long as the combined lengths of “the
pedicel and first funicle’ jomte.2 cee nies eee io eee ae pulla.
rg09] The Mymarid Genus Camptoptera. 29
LITERATURE REFERRED TO.
1856. Foerster, Arnold. Hymenopterologische Studien, Aachen, II, pp. 116,
119-120, 144.
1867. Kirchner, Leopold. Catalogus hymenopterorum Europae, Vindobonae,
p. 201, genus No. 784.
‘784. G. Camptoptera Foérst. (Pteroclisis Férst. Hym. Stud. II
144.)
1. C. pAPAVERIS Forst. Aachen. Aus Gallen V. Aulex rhoeadis.”’
1873. Snellen van Vollenhoven, S. C. Schetsen ten gebruike bij de Studie der
Hymenoptera, ’S Gravenhage (Uitgegeven door de Nederlandsche Ento-
mologische Vereeniging), 1869-1873, IV, Proctotrupiden, tabel XIII,
Camptoptera Forst.
Enlarged profile pen-drawing of the female Camptoptera (evidently the
species papaverts Foerster), with enlarged male antenna. The details are
incorrect for the female and as the work as a whole has been discredited,
it may be assumed that the details of the male antenna are as well non-
reliable.
1874. Kaltenbach, J. H. Die Pflanzenfeinde aus der Klasse der Insekten.
Stuttgart, p. 21.
1887. Cresson, Ezra Townsend. Synopsis of the Hymenoptera of America,
North of Mexico. Trans. American Ent. Society, Philadelphia, supplemen-
tary volume, 1887, pp. 85, 141. :
1889. Provancher, L’Abbe L. Additions et corrections au volume II de la faune
entomologique du Canada traitant des Hyménoptéres, p. 404. (Additions
et corrections a la faune Hyménoptérologique de la province de Québec.
November, 1888.) Jn Petite faune entomologique du Canada et particu-
liérement de la province de Quebec. Quatriéme ordre les Hyménopteres,
Québec, 1883.
1898. De Dalla Torre, Carl G. Catalogus Hymenopterorum hucusque descrip-
torum systematicus et synonymicus. Lipsiae, MDCCCXCVIII, V, p. 4381
and footnote.
1904. Ashmead, William Harris. Classification of the chalcid flies or the super-
family Chalcidoidea, with descriptions, etc. Memoirs Carnegie Museum,
Pittsburgh, I, No. 4 (Publications of the Carnegie Museum, Serial No. 21),
pp. 362, 369.
1905. Girault, Alecandre Arsene. Two new Mymaridae. Psyche, Boston,
Massachusetts, XII, p. 91.
1905. Stiles, Charles Wardell. The international code of zoological nomencla-
ture as applied to medicine. Bull. No. 24, Hygienic Laboratory, Public
Health and Marine—Hospital Service of the United States, Treasury Dept.,
Washington, D. C.
STUDIES ON APHIDIDA. II.*
Joun J. Davis.
OFFICE OF THE STATE ENTOMOLOGIST, URBANA, ILLINOIS.
The Birch Callipterus.
[Calli pterus betulecolens (Fitch) Monell.]
What is supposed to be this species was first described by Asa
Fitch in his “‘Catalogue of the Homoptera of New York” in 1851
as Aphis betulecolens. Monell described it from Missouri as
Callipterus betulecolens, n. sp., and placed Aphis betulecolens of
Fitch as doubtfully synonymous. Thomas hesitatingly placed
this species and Calaphts betulella as Callipterus betule of Koch,
but there is little doubt that all three are distinct species. Oest-
lund redescribed the winged viviparous female, from specimens
collected in Minnesota, as C. betulecolens (Fitch), and gave C.
betulecolens of Monnell as synoynmous. It has been further
reported in the literature by Miss Edith M. Patch from Maine,
and by W. T. Clarke from California.
In Illinois I have found it a very common species on Betula
alba, and it doubtless occurs wherever the birch is found. It
became very abundant this fall (1908), especially at the time the
sexual forms appeared.
The viviparous females were found on birch throughout the
summer and as late as September 28, at Chicago, Illinois. The
oviparous females, winged males, and eggs were first noticed
September 21, 1908, and at that time many pairs were observed
in copula. Careful observations were made to determine whether
or not the hind tibize of the oviparous females, which in this species
always bear sensoria, played any part in copulation or oviposition,
it having been stated by R. A. Vickery (‘‘A Comparative Study
of the External Anatomy of Plant Lice.’ p. 11. Separate of
the Twelfth Report of the State Entomologist of Minnesota, 1908)
that the hind tibiz of oviparous female aphids have some func-
tion in connection with oviposition. The hind tibiz of the
oviparous females of this species, however, played no part what-
ever in Oviposition, nor, so far as could be observed, in the act
* For No. 1 of this series see the Annals of the Entomological Society of
America, Vol. I, No. 4, December, 1908, pp. 251-264.
30
1909 | Studies on Aphididae IT 21
of copulation. During the period of oviposition the females
wander along the branches in search of suitable places on which
to deposit eggs. Upon finding such places—for example, a cre-
vice formed by a dormant bud or a small crotch where two small
branches or twigs unite—she extends the tip of the ovipositor
into the crevice and deposits the egg as far into it as possible.
This process requires from three to five minutes. Usually the
female seeks another part of the branch soon after laying the
egg, but seldom lays a second egg immediately. One individual
which had just oviposited was examined, and eight more well-
developed eggs were found in her body.
DESCRIPTIONS.
Winged viviparous female.—The description of the winged viviparous
female by Monell and Oestlund are quite complete, and further descriptions are
unnecessary except for the few remarks following: The basal two thirds of
antennal segment III bears 8 to 14 circular sensoria ina row. Usually the num-
ber is 12 or 13, and the extremes 8 and 14 are exceptions. Figure of the antenna
is given in plate V, figure 1.
Wingless oviparous female.—Head pale whitish yellow. Prothorax pale
greenish yellow with a longitudinal brownish stripe on each side. The remaining
thoracic segments like the first except that the lateral marking, which continues
on these segments, changes to a decided green. Abdomen yellowish with a
slight greenish tint, and sometimes the orange-colored eggs show through the
body, giving it an orange tint. The lateral thoracic stripes continue along the
sides of the abdomen, widening on one of the anterior segments, but continuing
posteriorly to the cornicles as a narrow stripe. Immediately anterior to the
cornicles is a transverse hand of a brownish green color, which connects the green
lateral markings. Posterior to the cornicles the abdomen is pale greenish yellow,
with no noticeable markings. Sometimes the green markings above described
are very indistinct, and the abdomen, in this case, is of a cream yellow color.
Head with slight frontal tubercles and a more or less conspicuous tubercle
on the front bearing the median ocellus, six moderately long capitate hairs pro-
jecting cephalad from the front, and one on each side of the dorsum of the head
at the base of the frontal tubercles. (PI. V, fig. 2.) Antennz longer than body,
usually about one and one half its length, segment III longest, being twice the
length of V, IV subequal with filament of VI, V about twice the length of
the base of VI, VI (base and filament) slightly less than III and the fila-
ment slightly more than twice the basal portion; distal halves of III and IV
with weak imbrication, distal half of V and all of VI with slightly stronger imbri-
cation, distal half of V and all of VI with slightly stronger imbrication; an
occasional minute capitate hair (Obj. 1-6) on the basal half of the antennz:
1 to 7 (usually 2 to 4) sensoria near the base of segment III. (PIL. V, fig. 3.) Seg-
ments I and II concolorous with head, III, 1V, V, and base of VI with the basal
halves whitish yellow and the distal halves blackish, filament of VI entirely black.
In older specimens the antenne are entirely black excepting I, II and a small por-
tion at basal end of each of the remaining segments. Beak not reaching beyond
the coxe of the second pair of legs. Eyesred. Legs with femora pale yellowish,
tibiae brownish excepting distal ends which are black, and the tarsi black. Hind
tibiae with the basal half noticeably swollen and bearing 75 ar 80 small, more or
less circular, and irregularly placed sensoria, most of which are borne ontthe
inner surface. (PI. V, fig. 4.) Abdomen with a transverse row of 6 or 8 tuber-
cles on each segment, each tubercle bearing a single moderately long capitate
hair. Cornicles typical of the genus, and of a pale greenish yellow color. (PI.
V, fig. 5.) Style concolorous with the cornicles, short and rounded at the apex
32 Annals Entomological Society of America [Vol. II,
(unlike the style of the viviparous female in that there is no constriction near
the base), and 12 to 15 hairs projecting caudad. (Pl. V, fig. 6.) Anal plate
larger than the style, similarly rounded, and with numerous hairs over its sur-
face. (This differs from the anal plate of the viviparous female in that it is not
divided or bifid.)
Measurements [taken from 10 specimens (Chicago, Ill, September 21, 1908)
mounted in balsam].—Length of body 2.00—2.54, av. 2.28 mm.; width, 0.82-1.21,
av. 0.97 mm.; antenna, I, 0.13; II, 0.065; III, 0.92-1.06, av. 0.98; LV, 0.54—0.67.
av. 0.60; V. 0.42-0.50, av. 0.47; VI, base, 0.23-0.27, av. 0.25; VI, filament, 0.52-
0.65; av. 0.61; total average, 3.105 mm.; cornicles, 0.126 mm.; style, 0.10 mm.;
hind tarsus, 0.16 mm.
Winged male:—The males are similar to the winged viviparous females but
they may be easily distinguished from them as follows: Antennal segment III
of male with one third to one half more sensoria than the equivalent segment of
the female antenna, and they extend the entire length of the segment in the for-
mer while in the latter these sensoria are found only on the basal two thirds.
Further, the abdominal segments of the male have short transverse bands on the
dorsum, these being absent on the abdomen of the winged viviparous female.
(Described from winged males collected 7m copula, at Chicago, Ill., Septem-
ber 21 and 28, 1908.) Head pale greenish with more or less of a brownish tint,
in a few cases dusky; with noticeable frontal antennal tubercles; the medial
ocellus prominent and projecting beyond the fore border. Antenne nearly bare;
24-28 transverse (usually a few are circular) sensoria in a row the entire length of
III, the usual large elongate sensorium at end of V and a similar one, with several
smaller sensoria bordering it, at distal end of the basal portion of VI; nearly twice
the length of the body, III the longest, being one third longer than IV, IV usually
slightly longer than filament of VI, often LV and filament of VI subequal and
occasionally the latter slightly the longer, V one half or slightly more than one
half of III, base of VI one third to one fourth the length of filament of VI and
together they are longer than IV but shorter than III. (Pl. V, fig. 7.) Seg-
ments I and II pale greenish, III dusky or brownish at base, the distal half dark-
ening to black, IV black excepting basal one third which is pale or dusky, V
and base of VI black except the basal thirds of each, which are whitish, filament
of VI black. Eyesred. Beak not reaching to coxe of second pair of legs. Pro-
thoracic segment pale greenish with an almost square brown marking on the
median dorsum. Thoracic shield dusky brown. Wings hyaline, basal two
thirds of the stigmal vein subobsolete, stigma pale, costal and subcostal veins
pale brownish, remaining veins blackish, and a small dusky brownish area at the
end of each vein except the costal and subcostal, and usually the first and second
anal. (Pl. V, fig. 8.) The first and second discoidals branching at about one
half the distance from the margin to the third discoidal. Legs with femora
pale brownish, tibiae black at extremities and brownish in the middle, tarsi entire-
ly black. Abdomen pale yellowish green with a median longitudinal black mark-
ing on the dorsum, consisting of short transverse bands ending on the antepenul-
timate segment. A longitudinal row of fine hairs on the dorsum extending along
the outer portion of the black patch (two hairs to each segment, one on each
side), a very small pale area at the base of each hair (in specimens mounted in
balsam). Cornicles and style pale yellowish green in color and characteristic
of this genus.
Measurements [from 10 specimens mounted in balsam, collected at Chicago,
Ill., September 21 and 28, 1908]:—Length of body, 1.36-1.86, av. 1.61 mm.;
width, 0.52—0.68, av. 0.57 mm. (at base of cornicles the average width is 0.46
mm.) ; length of wing, av. 3.00 mm.; width, av. 0.93 mm. ; wing expanse, approx.
6.50 mm.; antenna, I, 0.13; II, 0.065; III, 0.83-1.08, av. 0.93; IV, 0.56—-0.70,
av. 0.64; V, 0.44-0.54, av. 0.50; VI, base, 0.18-0.24, av. 0.21; V1, filament, 0.54—
0.69, av. 0.62; average total, 3.095 mm.; cornicles, 0.09 mm.; hind tarsus, 0.16
mm.
Egg :—Pale orange when first laid, and later changing to jet black.
1909 | Studies on Aphididae IT 22
The European Linden Callipterus.
[Callipterus (Eucallipterus) tilie (Linn.)]
This beautiful aphid I found to be common on the European
and American lindens in Chicago parks and in several nurseries
in the vicinity of that city. Ihave been unable to find any refer-
ence to this species as having been found in this country, but
Mr. J. T. Monell writes that he received specimens of this species
from Mr. Theo. Pergande, which were taken on linden at Wash-
ington, D. C., May 26, 1886.
This species was first described by Linnzus as Aphis tilie
and has since been placed in Callipterus by Koch, in Pterocallis
by Passerini, and recently in Eucallipterus by Schouteden. Ow-
ing to lack of hterature I have been unable to compare the syn-
onymy of this genus, and have therefore accepted that given by
Schouteden. Likewise the original description of the genus Pter-
ocallis is unavailable, but in Aphidide Italice, Passerini says
of the genus: ‘“‘Antennz articulo septimo breviore, ceterum ut
Myzocallis.””. The characters given for this genus by Buckton all
agree with those of the genus Callipterus, with the possible excep-
tion of the triangular fuscous patches at the ends of the veins.
The length of the filament of the sixth antennal segment, which is
used in separating Pterocallis, is an uncertain character, and: of
no generic value in itself, for the length of this portion of the sixth
segment may vary from an extremely long filament, as in Callip-
terus betulecolens, to one subequal to the basal portion as in C.
trifolit Mon.
Mr. Monell has called my attention to a note by Buckton which
illustrates the instability of the genus under discussion. Under
Pterocallis juglandicola, Buckton* says: ‘On account of the
size of the seventh antennal joint, possibly this species might be
ranged under Callipterus; I leave it, however, where Passerini
has placed it.”’ Although C. tive differs slightly from the typical
Callipterus, it does not seem sufficiently characterized for a sep-
arate genus, and I therefore retain the genus Callipterus with
FEucallipterus as a possible subgenus.
The habits of this species agree in every respect with those
of the genus Callipterus. This year (1908) the sexual forms were
found as early as September 15.
* Buckton, G. B. Monograph of the British Aphides, London, Vol. III,
1880, footnote, p. 33.
34 Annals Entomological Society of America [Vol DE
DESCRIPTIONS.
Winged viviparous female (from specimens collected in Chicago, Ill., July
23, 1908) :—Head pale yellow, with a blackish line on each side connecting with
the thoracic markings described later. (Pl. VI, fig. 19.) Antennz black ex-
cepting the whitish basal halves of IV, V, and basal VI, a short area at middle
of III, and the filament of VI dusky; 10 to 16 transverse sensoria on the basal
half of III, one circular sensorium near apex of V and another at the apex of the
basal half of VI, around which are several smaller sensoria. All segments, espec-
ially the distal ones, with strong imbrications which completely encircle the
antenne, giving them the annulated appearance common in the antenne of
Schizoneura, Tetraneura, etc. Entire length slightly more than that of the body,
III longest, it being one third longer than IV, which is slightly longer than, or
subequal to, V, VI (base and filament) very httle longer than IV, and the base
of VI somewhat larger than the flament. (Pl. V, fig. 9.) Eyes bright red.
Beak reaching slightly beyond the coxe of the first pair of legs. Prothorax pale
yellow and the remainder bright yellow. A very conspicuous black stripe ex-
tends along each side from in front of the eyes to the base of the wings. When
the wings are in a resting position the black costal area on the wings gives the
appearance of a continuous black lateral line the entire length of the body. The
mesothorax also bears a brownish longitudinal marking on each side of the me-
dian dorsal. Wings hyaline, the costal area from the base to the stigma brown-
ish to black, the stigma similarly colored excepting a patch on the outer margin,
and a brownish triangular patch at end of each vein; first and second discoidals
branching at about two fifths the distance from the tip of the wing to where the
third branches. (PI. V, fig. 10.) Two fore pairs of legs pale excepting the tarsi,
which are black. Hind pair pale excepting the black femora and tarsi, the
femora with about 15 or 20 pale circular areee which look very much like the sen-
soria of the antennze when seen through a low objective, but with a 1-6 objective
they are found to be of hypodermic origin. (Pl. V, fig. 11.) Abdomen pale
vellow, with two longitudinal rows of brown black spots on each side of the
median dorsal line, the row nearest the median line consisting of 8 spots and the
other row with 2 to 5 spots. Cornicles pale yellow with a fine, almost indis-
tinguishable, dark apical margin, tuberculate in shape, and about one third the
length of the hind tarsus. (PI. V, fig. 12.) Style concolorous with cornicles,
globular and often with the ‘‘neck’’ barely constricted; 15 or 20 fine tuberculate
hairs projecting caudad. (PI. V, fig. 13.) Anal plate decidedly bifid.
Measurements [taken from 12 specimens mounted in balsam]:—Length of
body, av., 1.97 mm.; width, av. 0.61 mm.; length of wing, av., 2.65 mm. ; width,
0.895 mm. ; wing expanse, approx., 6 mm.; antenna, I, 0.08; II, 0.065; III, 0.64—
0.74, av., 0.68; IV, 0.41-0.49, av., 0.44; V, 0.37-0.45, av. 0.41; VI, base, 0.26-—
0.32, av. 0.28; VI, filament, 0.195-0.245, av., 0.215; average total, 2.17 mm.;
cornicles, 0.057 mm.; style, 0.145 mm.; hind tarsus, 0.155 mm,
Wingless viviparous female :—I have never been able to find wingless vivi-
parous females, though I have made careful searches for them. Buckton has
described and figured what he calls the apterous viviparous female, but he does
not state when this individual was collected, and the characters given are too
incomplete to decide whether or not it is the viviparous, or whether it is the
oviparous. I believe he has described the oviparous female under the head of
apterous viviparous female, as his description compares exactly with the ovi-
parous females which I have found in Illinois.
Wingless oviparous female [from specimens collected in Chicago, Ill., Sep-
tember 15 and October 8, 1908]:—Head black or brownish, sometimes with a
pale median longitudinal line. (Pl. VI, fig. 20.) Antennal segments I and II,
dusky, III, IV, V, and base VI pale at basal halves or two thirds, and black
the remaining lengths, filament VI blackish; the usual sensorium at the end of
V, and one at end of the basal portion of VI, which is surrounded by several
smaller ones; imbrication slight on IV and V, but distinct on VI, although with-
out the conspicuous annulations of the winged viviparous antenne. (Pl. V,
fig. 14.) Total length slightly less than that of the body; III, the longest, being
1909] Studies on Aphididae II 25
slightly less than VI and about a third greater than IV; 1V and V subequal; basal
VI less than V and greater than filament VI. Eyes red. Beak not reaching or
barely reaching the coxe of the second -pair of legs. Prothoracic segment
brownish to black, with a small yellowish area on each side of the median,
and often with a threadlike pale median line. The coloration of the meso- and
the meta-thorax the same as that of the abdomen described below. The two
fore pairs of legs pale yellowish, excepting the dusky distal end of the tibia
and the black tarsi; hind pair with the tibia, and distal halves of the femora
dusky to black, and the tarsi black; hind tibize swollen and bearing about 35
irregularly placed, inconspicuous, circular sensoria on the basal halves. (Pl. V,
fig. 15.) The two posterior thoracic and the abdominal segments with a black
transverse band on each side of the median line, and a black spot on each side;
the interspaces yellow. The anterior dorsal bands necessarily shorter and the
bands on segments 5, 6 and 7 uniting in the middle, thus forming a single trans-
verse band on each of these segments. The position of the dorsal markings is
well illustrated by Koch in ‘‘ Die Pflanzenlause,’”’ figure 283. Cornicles blackish
and tuberculate as those of the viviparous female. Style pale yellowish. At
base of each cornicle and beneath the body, on each side, is a very noticeable
silvery white spot, which disappears on specimens mounted in balsam, the same
arece on the undersides of such specimens being represented by a brownish spot.
Measurements [from 8 specimens mounted in balsam]:—Length of body,
2:00 mm.; width, 0.89 mm.; antenna, I, 0.08; II, 0.065; III, 0.41—0.50, av.,
0:44: 1V, 0.28-0:30, av., 0.27; V, 0.24—0.30, av., 0.275; VI, base, 0.17-0.28, av.
0.20; VI, filament, 0.14—0.185, av. 0.165; average total, 1.495 mm.; style, 0.145
mm.; hind tarsus, 0.15 mm.
Winged male:—The male may readily be distinguished from the winged
viviparous female by its smaller size and by the sensoria on the antennae. The
antennez of the female bear sensoria only on the basal halves of III, and the usual
ones at the distal ends of V and base VI, while the male antenne bear them on
every segment excepting I and II. (Described from specimens collected at
Chicago, I1l., September 15 and October 8, 1908.) Head usually dark brownish
to blackish excepting a more or less triangular yellowish area at the posterior
portion; sometimes, however, the yellowish area extends to the fore border of
head. Antenne black excepting the pale dusky filament of VI and the extreme
base of all segments beyond III; annulations as on the antennz of the winged
viviparous female; total length a little more than that of the body, III the long-
est, and slightly less than twice IV, IV and V subequal, VI about two thirds III,
and the base VI longer than the filament; 30 to 38 transverse sensoria placed
more or less in a row the entire length of III, 6 to 9on IV, 7 to 9on V, 3 to 50n
base of VI, and the usual one at distal end of the same. (Pl. V, fig. 16.) Eyes
red. Beak not reaching the base of the coxz of the second pair of legs. Pro-
thorax yellowish with the typical black lateral stripe which extends from the
eyes to the base of the wings. Thoracic shield brownish to blackish. Wings as
those of the viviparous female, except that they are smaller. Fore pair of legs
yellowish or slightly dusky, with the tarsi black, hind pair with the femora,
bases of tibia, and tarsi blackish, and the middle pair usually intermediate
between these two types of coloration; the middle and hind femora usually with
the clear circular are referred to in the description of the viviparous female.
Abdomen pale yellowish green, with a row of short, transverse, blackish mark-
ings on either side of the median dorsal line, and a partial row of dots on each
side. Cornicles dusky and tuberculate as in the other forms. Style yellowish
or with a slight duskiness.
Measurements [taken from six specimens mounted in balsam]:—Length of
body, 1.42 mm.; width, 0.56 mm.; length of wing, 2.41 mm.; width, 0.87 mm. ;
expanse, approx., 5.50 mm.; antenna, I, 0.08; II, 0.065; III, 0,60—0.66, av.
0.68; IV, 0.34-0.37, av. 0.36; V, 0.32—-0.39, av. 0.385; VI, base, 0.21-0.26, av.
0.23; VI, filament, 0.14—0.21, av. 0.17; average total, 1.885 mm.; cornicles, 0.05
mm.; style, 0.10 mm.; hind tarsus, 0.13 mm.
30 Annals Entomological Society of America [Vol. iH
The Tulip-tree Aphid.
(Macrosiphum lirtodendri Monell, and var. rufa Monell.)
Macrosiphum lirtodendri is not an uncommon species in IIli-
nois, occurring on Liriodendron tulipifera, and has been observed
in the vicinity of Urbana for the past two years.
It was first described by Mr. J. T. Monell from Missouri, and
has since been reported from Washington, D. C., by Dr. L. O.
Howard, who noted that the eggs were common in the pupal shells
of Cicada which were attached to the tree trunks.
In Illinois, the eggs, which were first found the middle of Octo-
ber (1907), are laid in crevices, usually between the buds and the
branch. The following spring the eggs began to hatch between
March 29 and April 3, the young at this latter date being of the
first instar. The entire yearly cycle is passed on the tulip tree.
The variety rufa Monell was first observed July 6, inter-
mixed with the normal form in colonies and about equaling the
latter in abundance.
DESCRIPTIONS.
Winged viviparous female:—The original description by Monell* is suffi-
ciently complete except for a few minor points, but owing to the fact that this is
the only description published of the tulip tree aphid, and that it is inaccessible
to many workers, | will redescribe it from Mlinois specimens.
Head pale greenish yellow or dusky; with conspicuous frontal antennal
oe and the median ocellus projecting beyond the fore margin. (PI. VI
21.) Antenna black excepting the two basal segments, w hich are dusky
as 14-18, circular sensoria in a row on III, the usual conspicuous one at the
distal end of V, and one large one with sever ‘al smaller ones surrounding it at
the apex of base V I; more than one half longer than the body, III, IV and V
subequal, but II] alw ays slightly longer than either IV or Vie wel the longest,
being nearly twice V, base VI about one fourth the length of V, filament VI
long “and setaceous. (PL. VI, fig. 22.) Eyes black. Ocelli dark red and rather
conspicuous. Beak barely reaching the coxe of the second pair of legs. Thorax
brownish. Wings hyaline with dark brown veins, the first and second discoidals
branching at a distance less than one half from the margin to where the third
discoidal branches. (Pl. VI, fig. 23.) Legs black, excepting the basal four
fifths of the femora, which portion is pale greenish. Abdomen pale green.
Cornicles black excepting the extreme base, which is pale greenish; reaching
beyond the tip of the abdomen, and three to three and one half times the length
of the style; slightly incrassated at the base and barely noticeably dilated near
the apex; distal end distinctly imbricated. (Pl. V, fig. 17.) Style pale green-
ish yellow, ensiform, decidedly uncurved, and about one half longer than the
hind tarsus. (Pl. V, fig. 18.)
Measurements [irom five specimens mounted in balsam] -—Length of body,
2.25 mm.; width, 0.77 mm.; length of wing, 3.35 mm.; width, 1.18 mm.; ex-
panse, 7.50 mm.; antenna, I, 0. 135; TI, 0:07; III, 0:64— 0. 82, av. 0.74; LV, 0.66—
0:77, av. 0:705;'V, 0.70-0.74, av. 0. 715: Wilk. base, 0.175; VI, filament, 1.09-1.26,
av. 1.17; average total, 3.70 mm.; cornicles, 0.85 mm.; style, 0.26 mm.; hind
taruss, 0.16 mm.
* Notes on the Aplidide of the United States, with descriptions of species
occurring west of the Mississippi. U.S. Geol. and Geog. Survey of the Terri-
tories, V ‘ol. V, No. wot. (p20) taro,
1909] Studies on Aphididae II 37
Wingless viviparous female:—Head pale green and with frontal tubercles
as on winged. Antenna black excepting segments I and II, and a small basal
portion of III, which are pale green or slightly dusky; the usual sensoria on V
and VI (one exceptional specimen has 3 and 4 sensoria, respectively, on each
antenna near the base of III); total length about one half greater than that of
the body, segment III about one fourth longer than IV, 1V and V subequal, VI
the longest, and slightly less than IV and V together, base VI about one fourth
the length of V, filament VI setaceous and about one half longer than III (a
single exception, not recorded in measurements below, has III and filament VI
subequal). (PI. VI, fig. 24.) Eyes black. Beak reaching to or sligtly beyond
the coxe of the second pair of legs. Thorax pale green, with a very slight pul-
verulence. Legs pale green except the bases and apices of tibia, which areas are
dusky to black, and the tarsi which are black. Abdomen concolorous with, and
bearing a similar pulverulence as, the thorax. Ventral surface of body with the
whitish pulverulence or bloom more conspicuous. Cornicles black except the
bases, which are pale, similar in shape to those of the winged female, and about
two and a half times longer than the style. Style pale green, ensiform, and about
twice the length of the hind tarsus.
Measurements [from specimens mounted in balsam]:—Length of body,
ib —2250 mms width O17 2107 mm antenna TOs] Ee 0107. Wie 0:73); LV,
0.60; V, 0.61; VI, base, 0.16; VI, filament, 1.07; average total, 3.87 mm. ; cor-
nicles, 0.71 mm.; style, 0.29 mm.; hind tarsus, 0.15 mm.
Immature :—Dull green with blackish legs, antenne, cornicles, and eyes.
Wingless viviparous female, variety rufa:—I have found only the wingless
forms of this red variety. It is identical in form and measurements with the
normal form of the wingless viviparous female, and is only distinguished from
the latter by its coloration.
Head, thorax, and abdomen of a decidedly rosy tint. The ventral surface
pale reddish, and covered with a whitish pulverulence or bloom. Antennal seg-
ments I and II pale, III pale at base and gradually darkening to black, remain-
ing segments black. Eyes black. Coxe pale reddish, otherwise legs as in nor-
mal form. Cornicles black, excepting at bases which are pale. Style pale with
a reddish tint.
Wingless oviparous female:—The only structural character distinguishing
this form from the wingless viviparous female is the presence of numerous
sensoria on the hind tibiz. General color pale greenish, the abdomen with a
yellowish tint. Notes of the individual colorations of the body and appendages
have been lost or misplaced, but in general they agree with those of the wingless
viviparous female. The antenna is a little less than one and one half times the
length of the body, III slightly longer than IV, IV and V subequal, VI the long-
est, being subequal to IV and V together, base VI about one sixth or less the
length of the filament, which is one and one half times III; sensoria absent
excepting the usual ones at the distal ends of V and base VI. (PI. VI, fig. 25.)
The basal half of the hind tibize swollen and bearing 50 to 70 small circular sen-
soria. (Pl. VI, fig. 26.)
Measurements [from specimens in balsam and alcohol]:—Length of body,
1.97-2.47 mm.; width, 0.82-1.07 mm.; antenna, I, 0.125; II, 0.065; III, 0.52-
0.65, av. 0.59; IV, 0.42-0.59, av. 0.505: V. 0.42-0.59) av. 0.51; VI, base, 0.15; VI,
filament, 0.92—1.02, av. 0.95; average total, 2.89 mm.; cornicles, 0.60 mm. ; style,
0.22 mm.; hind tarsus, 0.145 mm.
Winged male [from specimens in balsam and _ alcohol]:—Distinguished
from the viviparous female by its smaller size and the numerous sensoria on
antennal segments III, IV and V.
Coloration similar to the winged viviparous female, excepting the usual
presence of a faint row of dusky to blackish dots on each side of the abdomen.
Head and appendages (excepting the antenne) similar in shape and markings
to those of the winged female. Antenne about one and one half times the length
of the body, III about one half VI, IV and V subequal, basal VI about one
fourth V, filament VI equal to IV and V combined; 40 to 53 circular sensoria
irregularly placed on one side of III, 22-27 more or less in a row on IV, 15-21
38 Annals Entomological Society of America [Vole il
xX
in a row and the usual larger one at distal end of V, a single large sensorium with
several smaller ones near its border, at distal end of base VI. (Pl. VI, fig. 27.)
Wings with the first and second discoidals branching slightly nearer to the mar-
gin than in the viviparous female, this character being more variable in the
latter.
Measurements [from specimens in alcohol and balsam]:—The measurements
for different individuals differs considerably in some cases, but the comparative
lengths are quite constant. Length of body, 1.79-2.36 mm.; width, 0.61—0.89
mm.; length of wing, 2.61—-3.29 mm.; width, 0.98—1.18 mm. ; expanse, 5.75—-6.50
mim. antenna, 1, 0111-11. 01072 TM. '0;56-0: 75; av, 0167- IV. (044=0'63) iave O53
V, 0.47-0.59, av. 0.535; VI, base, 0.18-0.145, av. 0.14; VI, filament, 0.99-1.26,
av. 1.12; average total, 3.175 mm.; cornicles, 0.55 mm.; style, 0.22 mm.; hind
tarsus, 0.15 mm.
Egg:—Pale greenish or greenish yellow when first laid, and later changing
to jet black. Elliptical oval, measuring 0.64 mm. in length and 0.31 mm, in
width.
The Ornamental Currant Macrosiphum.
(Macrosiphum cynosbatt Oestlund.)
The winged form of Macrostphum cynosbati was first described
by Professor Oestlund, it having been collected in Minnesota in
1887 on Ribes cynosbati. It has not since been reported in the
literature as occuring elsewhere.
This year (1908) it was very common and in some localities
quite injurious to the ornamental currants in Chicago parks.
It did not appear in injurious numbers until the first of August,
at which time only the wingless forms were present, and these
densely colonized the tender terminal twigs, infesting all parts,
but particularly tips of the twigs, or, when present in abundance,
the under sides of the leaves also, and in sufficient numbers to
curl them more or less. Winged forms were first found in these
colonies September 3. Usually either the common brown ant
(Lasius niger var. americanus) or the black ant (Formica fusca)
was in attendance upon the plant-lice.
The sexual forms of this species were first found on these
same bushes October 5, at which time they were observed pairing.
DESCRIPTIONS.
Winged viviparous female [from specimens collected in Chicago, Ill., Aug-
ust 10, and September 3 and 11, 1908]:—Head dusky with distinct and approxi-
mate frontal antennal tubercles and 6 or 8 subcapitate hairs projecting cephalad
from the fore border. Antenne black, excepting the two dusky basal segments,
and a fine pale ring on III at joint of II and III; sparsely covered with hairs
which are more or less subcapitate; 30 to 43 tuberculate, circular sensoria, irreg-
ular in size and arrangement, placed the entire length of III, occasionally a single
one on IV, the usual sensorium at the distal end of V and one at apical end of
base VI with about 6 smaller ones beside it; total length about one half longer
than the body, III almost twice longer than IV and a little more than one
fourth shorter than VI, IV and V, subequal, IV usually very slightly the longer,
base VI very short, it being subequal to I, filament VI about three times V.
(Pl. VII, fig. 30.) Eyes dark red. Beak reaching slightly beyond the coxe
of the hind pair of legs. Thorax pale green excepting the shield, which is pale
1909] Studies on Aphididae II 39
brownish, Wings hyaline, with brownish veins, the first and second discoidals
branching at one half or less the distance from the margin to where the third
discoidal branches. (Pl. VII, fig. 31.) Legs pale green to the distal portion of
femur, which is blackish, tibia pale brownish excepting the black apical end,
tarsus black. Abdomen entirely pale bright greenish. Cornicles pale greenish,
white at basal half, the distal portion changing to blackish; reaching beyond
the tip of the abdomen and more than three times the length of the hind tarsus;
cylindrical and slightly dilated at distal one third. (Pl. VI, fig. 28.) Style
concolorous with abdomen, ensiform, and slightly less than one-half the length
of the cornicles. (Pl. VI, fig. 29.)
Measurements [from alcoholic and balsam mounts, collected in Chicago,
September 3 and 10, 1908]:—Length of body, 1.65 mm.; width, 0.61 mm,; length
of wing, 2.65 mm.; width, 0.91 mm.; expanse, approx. 6.00 mm.; antenna, I,
0.08; II, 0.07; III, 0.55-0.77, av. 0.65; IV, 0.32-0.42,. av. 0.35; V, 0.29-0.36, av.
0.32; VI, base, 0.08—-0.09, av. 0.085; VI, filament, 0.83-1.09; av. 0.95; average
total, 2.505 mm.; cornicles, 0.48 mm.; style, 0.20 mm.; hind tarsus, 0.14 mm.
Wingless viviparous female:—Head with about six subcapitate hairs pro-
jecting cephalad from the front. (Pl. VII, fig. 32.) Antenne borne on the
usual frontal tubercles; pale excepting segment VI and the distal portion of V;
usually 15 to 26 circular sensoria, unequal in size and irregularly placed on III
(this is a very uncertain character in this form, some antenne bearing only 7
sensoria on III), and the usual ones at apical ends of V and base VI; longer than
body, the comparative lengths of the separate segments as given for the winged
viviparous female. (Pl. VII, fig. 33.) Eyes dark red. Beak reaching or
extending slightly beyond the coxe of the third pair of legs. Legs pale, except-
ing the distal ends of the tibia, and at the joints of the tibie and femora, which
are dusky, and the tarsi which are black. Abdomen bright green. Cornicles
pale excepting the tips, which are slightly dusky, and about four times the
length of the hind tarsi. Style pale with a slight greenish tint, and about two
fifths the length of the cornicles.
Measurements [from specimens mounted in balsam]:—Length of body,
190 mime width, O.88mm:;antenna, I, 0/095; Tl, 0:08; TIl, 01615 1V, 0:35. V,
0.35; VI, base, 0.095; VI, filament, 0.89; average total, 2.47 mm.; cornicles,
0.61 mm.; style, 0.23 mm.; hind tarsus, 0.15 mm.
Wingless oviparous female:—Head pale yellowish or greenish yellow, bear-
ing about six subcapitate hairs which project cephalad from the fore margin.
Antennal segments I and II concolorous with the head or slightly dusky, the re-
maining segments dark to black; subcapitate hairs sparsely placed on all seg-
ments; 13 to 23 circular sensoria irregularly placed on III, and the usual ones
at the apical ends of V and base VI: subequal to or slightly longer than the
body, III one third longer than IV, IV and V subequal, VI the longest, being
about three times V, the base VI about subequal tol. (Pl. VII, fig. 34.) Eyes
dark red. Thorax pale yellowish. Legs pale excepting the distal ends of the
tibiz and femora, also the tarsi, which are blackish; the basal one third of the
hind tibiz swollen and bearing 30 or 40 irregularly arranged circular sensoria,
placed on the inner surface. (PI. VII, fig. 35.) Abdomen pale orange and
usually with two or three yellowish spots which are the eggs showing through
the semi-transparent body tissue. Cornicles pale excepting at tips, which are
dusky, cylindrical, and sometimes slightly dilated at the distal third, subequal
in length to segment III of the antenna. Style pale and about two fifths the
length of the cornicles.
Maesurements [specimens mounted in balsam]—Length of body, 2.09 mm. ;
width, 0.92 mm.; antenna, I, 0.11; I], 0.08; III, 0.60; IV, 0.37; V, 0.33; VI, base
0.11; VI, filament, 0.88; average total, 2.48 mm. (An exceptional specimen
had the following antennal measurements: III, 0.39; 1V, 0.26; V, 0.26; VI,
base, 0.08; VI, filament, 0.61 mm.); cornicles, 0.56 mm.; style, 0.28 mm.; hind
tarsus, 0.15 mm.
Winged male:—Head brownish and with very small subcapitate hairs
arranged as on wingless oviparous female. Antenna dark brown at the base,
darkening to black at the apical portion; bearing subcapitate hairs which are
less conspicuous than those on the antenna of the above described female; 20 to
40 Annals Entomological Society of America EV oleh
35 (usually 30 to 35) circular sensoria, irregularly placed on III, 11 or 12 on IV,
and 4 to 7 on V, with the usual larger distal ones on V and base VI; about one
third longer than the body, III more than one half longer than IV, VI and V sub-
equal but IV usually slightly the longer, VI the longest, it being about one half
longer than the next longest segment (III), base VI subequal with I. (Pl. VII,
fig. 36.) Eyes dark red. Beak reaching slightly beyond the coxe of the third
pair of legs. Thorax brownish. Wings as described for the viviparous female.
Legs with femora pale greenish excepting the distal ends, tibia brownish except-
ing distal ends and tarsi black. Abdomen blackish (no color notes taken). Cor-
nicles as in the other forms, the basal half pale greenish and the apical portion
dusky to black. Style pale brownish.
Measurements [from specimens in balsam]:—Length of body, 1.61 mm.:
width; 0.62; length of wing, 2.40 mm.; width, 0.86 mm.; expanse, approx. 5.25
mm.; antenna, 1, 0:09; Il, 0.075; IIT, 0:56; IV, 0:33: V., 0:28; V1, base, 0:08:
VI, filament, 0.80; average total, 2.215 mm.; cornicles, 0.40 mm.
The Button Bush Aphis.
(A phis cephalantht Thomas.)
This species was first described by Thomas in 1877 from speci-
mens collected in southern Illinois on Cephalanthus occidentalis.
The only addition to the accounts of its distribution or life history
is a single record by Mr. J. G. Needham* of its occurence on the
button-bush at Lake Forest, I.
I have found it very abundant on the same species of shrub
at Chicago, Hl., and Mr. J. T. Monell has kindly sent me authentic
specimens collected by him at St. Louis, Mo. September 29, 1908,
at Chicago, where the branches, especially the terminal shoots of
the button-bush were completely covered with this aphid, and
numerous eggs, which had been deposited without any seeming
care as to location, were found on the branches.
DESCRIPTIONS.
Winged viviparous female:—Head black, rounded at frontal margin, and
sometimes the median ocellus projecting slightly beyond the front. (PI. VII,
fig. 37.) Antenna blackish or black; 10-17 ‘circular sensoria irregularly placed
on III, usually 1-6 on IV, the usual one at the distal end of V and. a single large
one with several smaller ones about its margin, at distal end of base VI: total
length about three fourths that of the body, III about one half greater than
IV, 1V and V subequal, VI the longest, being subequal to IV and V combined,
base VI about one third III and filament VI subequal to III but invariably very
slightly longer. (Pl. VII, fig. 38.) Eyes blackish with a barely perceptible
reddish tint. Beak nearly reaching the coxe of the hind pair of legs. Thorax
black. Wings hyaline, with pale veins, the first and second discoidals branch-
ing at a point varying from one third to nearly one half the distance from the
margin to where the third discoidal branches. (Pl. VII, fig. 39.) Legs pale
brownish except at the joint of the tibia and femur, distal end of the tibia and
the tarsus, all of which are black; the hind pair of legs usually has the distal half of
the femur likewise black. Abdomen dark greenish brown to reddish brown; bear-
ing a row of white pulverulent spots on each lateral margin; in well prepared
balsam mounts a fleshy tubercle is seen on each side of each of the segments,
anterior to that bearing the cornicles and likewise on the segment posterior to
it; and a dark spot at base of each cornicle. Cornicles black, cylindrical, about
one half longer than the style, and not reaching the end of the abdomen. (PI.
* Psyche, Vol. 10, Feb. 1903, pp. 26; 27.
1909 | Studies on Aphididae IT AI
VII, fig. 40.) Style black, conical, an slightly longer than the hind tarsus.
Cele Wall fie.)
Measurements [from specimens in balsam]:—Length of body, 1.50-1.97
mm.; width, 0.70-1.14. mm.; length of wing, 2.37 mm.; width, 0.90 mm.;
expanse, approx. 5.25 min.; antenna, I, 0.065; II, 0.06; III, 0.30-0.34, av. 0.31;
IV, 0.195-0.235, av. 0.22; V, 0.195-0.22, av. 0.205; VI, base, 0.095-0.115, av.
0.105; VI, filament, 0.29—-0.365, av. 0.34; average total, 1.305 mm.; cornicles,
0.23 mm.; style, 0.15 mm.; hind tarsus, 0.1385 mm.
Wingless viviparous female:—Entire body pulverulent. Head dark red-
dish brown. Antennz blackish, comparative lengths similar to those of the
winged viviparous female, the usual sensoria at distal ends of V and base VI.
(Pl. VII, fig. 42.) Eyes blackish. Thorax concolorous with head. Legs pale
brownish, distal third of the femora, end of tibie, and the tarsi black. Abdo-
men brownish red, with a dusky marking at base of each cornicle which some-
times extends across the abdomen. Cornicles and style blackish.
Measurements [from specimens in balsam]:—Length of body, 1.29 (shriveled
—1.79 mm.; width, 0.68 (shriveled)—1.00 mm.; antenna, averages, I, 0.065;
II, 0.06; III, 0.30; IV, 0.20; V, 0.17; VI, base, 0.10; VI, filament, 0.33; average
total, 1.22 mm.; cornicles, 0.21 mm.; hind tarsus, 0.14 mm.
Wingless oviparous female:—Entire body more or less covered with a
whitish pulverulence which is densest at the posterior end. In most specimens
the pulverulence appears more distinct in four longitudinal rows of short trans-
verse areas on the dorsum. Head blackish or blackish brown, darker than the
thorax and abdomen. Antennae with segments I and II dusky; III pale brown-
ish, IV darker, V and VI blackish; about one half the length of the body, III
about one fourth longer than IV, IV and V subequal, VI the longest, it being
nearly twice as long as III, base VI about one third shorter than V, filament VI
subequal to IV and V combined; the usual sensoria at distal ends of V and base
VI. (PI. VII, fig. 43.) Eyes black. Thorax reddish brown to blackish brown.
Legs brown, excepting distal ends of the femora and tibie, and the tarsi, which
are blackish; hind tibiz much swollen almost their entire lengths and each bear-
ing approximately 80 more or less circular sensoria. (Pl. VII, fig. 44.) Abdomen
concolorous with thorax and with a small blackish area at base of each cornicle;
and fleshy tubercles along the sides as on the winged viviparous female. Cor-
nicles black and nearly one and one half times the length of the hind tarsi.
Style blackish and subequal in length to the hind tarsus.
Measurements [from specimens in balsam and alcohol]:—Length of body,
1.79-1.97 mm.; width, 0.93-1.00 mm.; antenna, averages, I, 0.05; II, 0.05; III,
0.20; IV, 0.165; V, 0.16; VI, base, 0.10; VI, filament, 0.27; average total, 0.995
mm.; cornicles, 0.19 mm.; style, 0.14 mm.; hind tarsus, 0.14 mm.
Winged male:—Readily distinguished from the viviparous female by its
smaller size and the presence of several sensoria on segment V of the antenna.
Head black and similar in shape to those of the other forms. Antennz
black; 14-18 circular sensoria irregularly placed on III, 9-12 on IV, 4-7 and the
larger distal one on V, and the usual one, with several smaller ones bordering
it, at distal end of base VI; about one half the length of the body, III one third
to one fourth longer than IV, IV and V subequal, VI about twice the length of
III, base VI one third shorter than V, filament VI about twice V. (Pl. VII, fig.
45.) Eyes blackish. Thorax black. Wings hyaline, the first and second dis-
coidals branching at two fifths to one third the distance from the margin to
where the third discoidal branches. Legs blackish excepting the basal halves
of the femora. Abdomen brownish to reddish brown, a dusky to blackish patch
at the base of each cornicle. Cornicles and style black.
Measurements [from specimens in balsam and alcohol]:—Length of body,
1.18-1.50 mm.; width, 0.465-0.54 mm.; length of wing, 1.97 mm.; width, 0.76
mm.; expanse, approx., 4.50 mm.; antenna, averages, I, 0.05; II, 0.05- III, 0.21;
IV, 0.155; V, 0.15; VI, base, 0.10; VI, filament, 0.29; average total, 1.005 mm.;
cornicles, 0.12 mm.; style, 0.10 mm.; hind tarsus, 0.13 mm,
Egg:—Yellowish brown when first laid, changing to jet black within a day
or so; elliptical-oval in shape, and measuring 0.31 mm. in width and 0.64 mm.
in length.
42 Annals Entomological Society of America [Vol It,
EXPLANATION OF PLATES V, VI, VII.
Callipterus betulecolens (Fitch) Monell.—Plate V, figs. 1-8.
Figure 1, antenna of winged viviparous @ ; 2, head; 3, antenna; 4, hind
tibia; 5, cornicle; 6, style of wingless oviparous Q ;7, antenna; 8, wing of winged
G..
Callipterus (Eucallipterus) tiie (Linn.).—Plate V, figs. 9-16; Plate VI,
figs. 19-20.
Figure 9, antenna; 10, wing; 11, hind femora; 12, cornicle; 13, style of
winged viviparous 9 ; 14, antenna; 15, hind tibia of wingless oviparous Q ; 16,
antenna of winged ¢; 19, head of winged viviparous Q ; 20, of wingless ovipar-
ous 9.
Macrosiphum lirtodendrit Monell.—Plate V, figs. 17-18; Plate VI, figs. 21-27.
Figure 17, cornicle; 18, style; 21, head; 22, antenna; 23, wing of winged
viviparous 9 ; 24, antenna of wingless viviparous 2 ; 25, antenna; 26, hind tibia
of wingless oviparous Q ; 27, antenna of winged <.
Macrosiphum cynosbatt Oestlund.—Plate VI, figs. 28-29; Plate VII, figs.
30-36.
Figure 28, style; 29 cornicle; 30, antenna; 31, wing of winged viviparous 9 ;
32, head; 33, antenna of wingless viviparous @ ; 34, antenna; 35, hind tibia of
wingless oviparous 9 ; 36, antenna of winged <.
Aphis cephalantht Thomas.—Plate VII, figs. 37-45.
Figure 37, head; 38, antenna; 39, wing; 40, cornicle; 41, style of winged
viviparous 2 ; 42, antenna of wingless viviparous Q ; 43, antenna; 44, hind tibia
Q
of wingless oviparous @ ; 45, antenna of winged .
ANNALS E. S. A. VOL: Il, PEATE ¥:
J. J. Davis.
ANNALS E. S. A. VoL. II, PLATE VI.
J. J. Davis.
Vou. II, PLATE VII.
ANNALS E. S. A.
a
J. J. Davis.
THE TRACHEAL SUPPLY IN THE CENTRAL NERVOUS
SYSTEM OF THE LARVA OF CORYDALIS CORNUTA L.
Witiiam A. Hitton, Cornell University.
This study was begun with two main objects in view. First,
to determine some land-marks for use in a study of the finer
structure of the ganglia and nerve cells of this insect; and second,
to determine the exact air supply of each ganglion for the pur-
poses of later experimentation.
It is well known that all of the organs of insects are abun-
dantly supplied with tracheae and many fine tracheoles, but the
number and dense arrangement of the minute tracheoles in many
of the organs comes as a surprise to the person who sees them
under a microscope for the first time. The central nervous sys-
tem is no exception to other organs, indeed the tracheoles are
very abundant in all parts of the nervous system.
One of the disputed points in regard to the tracheae is the ques-
tion whether the minute tracheoles anastomose with each other
or not. They are very abundant, they branch to some degree
and they are very long and wind and twist themselves into such
a dense mat that it is very hard to follow individuals and hard
to tell whether they unite or not.
There seems to be very little literature on the tracheal system
of Corydalis. N. Young Burke, in an unpublished thesis in the
library of the Department of Entomology of Cornell University,
shows the main tracheae of the tracheal system of Corydalis. In
her figures, ventral transverse tracheal branches running from
side to side are shown. It is from these ventral transverse branch-
es that most of the ganglia are supplied.
The subesophageal ganglion has two branches which come
from the transverse ventral cephalic trachea and run to the ceph-
alic part of the ganglion on either side. This ganglion is also
supplied by two other branches, one on either side in its caudal
region. These come from another source more caudad of the
ganglion.
The three thoracic ganglia all receive two branches, one on
each side from a ventral transverse trachea, which like the
tracheal branch of the upper part of the subesophageal ganglion
come to the ganglia from a cephalic direction.
46
1909] Trachael Supply, Nervous System Corydalis 47
The first to the seventh abdominal ganglia are supplied by one
tracheal branch on each side, but these come to the ganglia from
a caudal direction because the transverse tracheae in every case
cross below or caudal to the ganglion to which they send branches.
The transverse tracheae of the abdominal region except the first,
are smaller as a rule than those of the thoracic region and the
branches which run to the abdominal ganglia are often of consid-
erable length.
The eighth abdominal ganglion is supplied from the outside
by two long tracheal branches which come to it from the caudal
direction.
The brain as described by Krauss ’84, was found to be sup-
plied by three lateral branches on each side from a common trunk.
The following pages will give a more or less detailed descrip-
tion of the tracheal supply in each of the ganglia of Corydalis.
For the larger branches ordinary dissection under a lens was
the method of study. For finer details the entire central nervous
system was removed and mounted in glycerine to which a drop of
formalin was added. These mounts cleared up in a short time,
the air in the finest tracheal tubes remained and they could easily
be followed. No sections were studied at this time, but later, a
large series of preparations will be examined with several objects
in- view, among them the determination if possible of how the
finest tracheoles end.
THE BRAIN.
The brain is supplied by three main branches on each side, all
of which come from a large cephalic tracheal branch. Thesethree
seem to be lateral portions from the large common trunk which
runs forward somewhat reduced to supply other parts of the
head.
The three trunks on each side, furnish each lateral half with
all of its main tracheal branches. (Fig. A and 2.) There are
comparatively few tracheae that cross the middle line and most
of these are small. These three main portions enter the brain
by their branches laterally from the ventral side and the des-
cription of them will be given as though they were seen from the
ventral side.
The most cephalic part is the smallest and the most caudal
the largest as a rule. Each part has both superficial and deep
branches, and although the territories of the divisions overlap to
some degree, the areas supplied by each are somewhat as follows:
48 Annals Entomological Society of America (Wel GLE
(1) Cephalic branch, dorsal and lateral portion including
branches that go to the optic nerves. It divides into two main
parts before entering the brain, a cephalic smaller branch which
divides into a number of long tracheae, many of them superficial
and supplying the latero-cephalic portion of the brain. The larger
more caudal part of four or more rather large branches, for the
most part deep ones which supply the central part of the brain
and to some extent the other side.
(2) The middle branch, supplies deep parts to some degree,
particularly in the latero-caudal region near the connective, includ-
ing a large trunk which runs down it nearly to the subesopha-
geal ganglion. On entering the brain this large middle branch
divides into a number of deep portions.
(3) The caudal branch, gives superficial tracheae to the whole
mesal half of the brainand deeper ones which penetrate the ventral
side of the ganglion near the middle line at the caudal edge. It
is of two main parts, the shorter portion which divides again soon
into a number of branches near the middle part of the brain and
a longer more ventral portion. Each of these main branches on
running to the ventral side of the brain divides into tracheoles
before entering, and the larger superficial, are easily seen from the
ventral side, while the larger deep branches can often only be seen
as they turn in to send deep tracheoles to the central part of the
brain and smaller superficial branches to the dorsal side. The
large superficial ventral branches send off many fine superficial
tracheoles which may be seen over all parts of the brain, in some
degree parallel to larger when larger branches are near.
In the center of the brain on each side, there was found a much
darker area due to the mass of tracheoles which formed a central
dense tuft. (Figs. A, G, & 1 & 2.)
Following the arched nerve on each side there is a small tra-
cheal branch from the brain which supplies the small frontal gan-
glion, while on each side from some other lateral source a minute
lateral branch also supplies this ganglion.
THE SUBESOPHAGEAL GANGLION.
The two pairs of branches supplying the ganglion are from
two sources:
First, a ventral transverse tracheal branch of large size which
runs across under the ganglion and a little above it. It sends off
on each side a rather large tracheole which enters the ganglion
at its cephalic and ventral surface.
1909] Trachael Supply, Nervous System Corydalis 49
Second, two large branches which come from large lateral
tracheae and run up to the lateral and ventral caudal edges of the
ganglion,
The first, or more cephalic of these pairs of tracheae give off
branches to the upper part of the ganglion. Upon entering, each
divides at once into about two main parts and these short branch-
es divide again into several and branches from these supply by
both superficial and deep portions, the upper half of the ganglion,
with a tracheole for each connective with the brain and small
tracheoles to each of the nerves which come off in this region.
The branches which go to the nerves and the connectives break
up into a very fine network of tracheoles without anastomosis.
In the case of the connectives, the area supplied is less than one
half the length of the distance to the brain. The fine tracheoles
in the connectives and the larger nerves run in a sort of circular
or spiral manner. Many minute tracheoles are given off from all
the branches in the ganglion.
The second or caudal pair of tracheae are larger than those
just described. They come from long lateral branches from below.
They supply the lower half of the ganglion, including the con-
nectives, with tracheae and tracheoles. On entering the ganglion
they each divide at once into about two main parts; the more
cephalic portion of a number of branches which run towards the
cephalic part of the ganglion. The other smaller portion consist-
ing chiefly of a large branch which supplies the most caudal part
of the ganglion with small tracheae and a large branch which sup-
plies the connective of its own side by one very long twig and
others shorter.
Superficial and deep branches form a dense network through-
out the ganglion. There seem to be about as many small trach-
eoles in the middle line as at the periphery. Branches from one
side cross to the opposite side in considerable numbers.
Branches from the upper and lower supplying trachae interlace
considerably. There are many long fine tubules with few side
branches, few free ends, and no anastomoses. Although the trach-
eoles are abundant in all parts of the ganglion, the densest mass
of them is in the central part, on each side.
The longest connective branches from the lower tracheae run
only about one third of the length of the connectives between this
ganglion and the first thoracic. The connectives between these
two ganglia are very long and their middle portion is supplied on
50 Annals Entomological Society of America [Vol. II,
each side by a tracheole not connected directly with the tracheal
system of either, but from an outside source. The tracheal sys-
tem of the connectives is very complete, many fine long branches
wind and interlace about the connectives with few free ends and
no anastomosis, although the long fine branches of different sys-
tems mingle very intimaetly. (Fig. B, H & 3.)
FIRST THORACIC.
This ganglion is suppled by a transverse branch above it
much like that supplying the cephalic part of the previously des-
cribed ganglion. From this transverse branch or rather arch,
branches one on each side enter the ganglion laterally and a little
toward the head end. ‘There are two main branches given off
at once on each side. On the left side the more cephalic is in
large part a connective branch, which divides into two at the
cephalic part of the ganglion; the outer of these runs up the out-
side of the connective for a long distance and its branches mingle
with those of the median independent branch of the connective.
The inner of the two just mentioned, runs to the point of origin
of the cephalic connectives and sends a somewhat smaller branch
along the inner side of each connective as well as a few short
tracheoles which turn back into the ganglion, between the con-
nectives. Both of these inner and outer connective branches
send a number of supplying tracheoles to the upper third of the
ganglion. On the left side the more caudal portion runs into and
along the edge of the ganglion, towards the caudal connectives,
distributing branches to the lower part of the ganglion and send-
ing a few short branches to the connective of its own side. On
the right side the larger more cephalic branch is quite different.
Similar parts of the ganglion are supplied by it as by the most
cephalic branch of the left side, only there seem to be more branch-
es and there is only one small connective twig which runs on the
outer side of the connective of the right side. The more caudal
branch of the right side is more like its fellow of the opposite side
in the number and position of the branches. In addition, a large
recurrent portion was traced way up almost to the origin of the
cephalic connective of the same side.
The above is a description of the main branches of the first
thoracic in a single specimen. Variations of this were found in
other specimens, particularly in the cephalic and caudal regions
of the ganglion in relation to the connectives. The supply of the
Tg09g| Trachael Supply, Nervous System Corydalis 51
cephalic connectives was more often nearly symmetrical and in a
number of specimens there was an additional median supplying
branch to the ventral region of the ganglion. This median sup-
plying branch ran under the ventral side of the ganglion, or a
little distance caudad of it on the connectives. It was found to
be fully as large as one of the branches of the lateral supply. In
about half of the specimens this branch was found and was appar-
ently torn off in one or two of the other specimens. This un-
paired branch divides into three a little below the ganglion, the
two lateral tracheae run up to the base of the ganglion, one on the
dorsal side, one on the ventral side and each breaks up into num-
erous tracheoles about the base of the ganglion near the middle
line. The median branch runs also in a cephalic direction for a
short distance and then divides into two and these again into
two main parts on each side, the more cephalic portion bends out
and after breaking into a number of smaller tracheae, supplies
the more caudal and lateral parts of the ganglion, while the more
caudal portion bends back with few twigs to supply the connective.
(Fig. 4.)
The various larger branches mentioned in connection with
this ganglion each give off an intimate supply of fine tracheoles,
both superficial and deep and although a certain area has more
tracheoles from a particular trunk, there are also numerous
tracheoles winding in all directions from other systems. The area
within the ganglion which receives the best supply of fine trach-
eoles is the central part on each side. The median line is rather
clear because there are few branches in this region, although a
number of the tracheoles from both sides cross this area in every
direction. (Fig. 4.)
SECOND THORACIC.
In a similar way to the first ganglion a branch on each side
enters from the transverse abdominal tracheal trunk. These two
branches run down laterally and to the ventral and slightly
cephalic region of the ganglion.
On each side the supplying trunk divides into three main
portions. A cephalic, consisting of a single long branch to the
outer side of the connective and a small tracheole from this which
runs more medially on the cephalic connectives. A median part
which breaks up into about five branches to supply the central
and cephalic part of the ganglion and a caudal branch. This
caudal portion is much smaller on the left side and furnishes small
Annals Entomological Society of America [Vol. II,
OL
to
tracheoles to the internal, lateral and caudal portions of the gan-
glion, but not much to the connective. The right corresponding
branch, however, is large and besides supplying tracheae to the
ganglion gives off two main parts which supply the caudal con-
nectives.
Variations from the above description were found in other
individuals studied. In some the supply to the ventral connec-
tives were symmetrical, or chiefly from the left side. The supply
of the cephalic connectives although more uniform was some-
times less symmetrical. And the cephalic connectives were often
supphed at their origin by branches from a number of systems.
(ign Fand-5%)
As in previous ganglia the nerves were each supplied by
usually one small branch, each from the nearest tracheal system.
The distribution of the smaller tracheoles was much like that
in the first thoracic ganglion.
THIRD THORACIC.
The tracheal supply is much the same in this as in the pre-
vious ganglion.
In the specimen of the series most described as a basis for the
larger branches, the left hand caudal branch instead of the right
hand one furnishes most of the connective supply.. (Fig. D, and
6.)
FIRST ABDOMINAL.
In the thoracic ganglia, the ventral transverse tracheal
branch which sends out two tracheae to each ganglion crosses
above or cephalic to the ganglia. The transverse tracheal
branches supplying the next seven, cross caudal to the ganglia
which they supply.
The first abdominal ganglion is much like the others in its
main branches. In the series most described and figured the
cephalic connective supply is symmetrical, the caudal supply
mostly from the left. In other specimens mostly from the right
with now and then a slightly less symmetrical distribution in the
cephalic region. This ganglion and the following show a more
definite lateral localization of the fine branches. (Fig. D and 7.)
SECOND ABDOMINAL.
Similar to the first. The main trunks come to it more medi-
ally. On the right side of the one of the series, the lower or cau-
dal connective is the larger. (Fig. 8.)
1909] Trachael Supply, Nervous System Corydalts 5
ies)
THIRD ABDOMINAL.
Similar. More medial supply. Quite a symmetrical distribu-
tion of branches in all specimens studied. (Fig. 9.)
FOURTH ABDOMINAL.
Very similar to the last. In the type specimen there is an
additional inner supply to the cephalic connectives from the left.
(Fig. ro.)
FIFTH ABDOMINAL.
Similar to the others. In the specimen described, the left
cephalic connective mostly supplied from the left side. The cau-
dalfitommthe tight. (Fig. 11.)
SIXTH ABDOMINAL.
Nearly symmetrical in regard to the cephalic and median
branches in the type, although the left is a little larger. The
caudal connectives supplied largely from the left. (Fig. 12).
SEVENTH AND EIGHTH ABDOMINAL.
The seventh ganglion has quite a symmetrical distribution.
The most cephalic branches on either side run up the connectives,
each on its own side. Two main central parts give tracheoles to
the main part of the ganglion, while a very large caudal part on
each side runs down the connective with a few short branches to
it and then breaks up into two main parts to supply the upper
portion of the eighth ganglion, although branches run way down
to the very tip of the ganglion as well. Two other main trunks
come to the eighth. Two rather large tracheae run up from the
caudal and lateral tracheal trunks to enter the ganglion ventrally
and laterally. These two last, differ from the other supply of the
other ganglia in not being connected with a ventral transverse
tracheal branch. Each of these last mentioned breaks up into
four or five main trunks and these supply the lower two thirds of
the ganglion as well as the numerous nerves which take their
origin from this region. (Fig. F and 13.)
The conditions just described for the Eighth abdominal gan-
glion are very uniform.
As may be seen from the description of the individual ganglia
of larval Corydalis, special branches are the only ones which
supply the central nervous system with air. These special
branches are three on each side to the brain from a large cephalic
branch; two pairs to the subesophageal ganglion, the cephalic
54 Annals Entomological Society of America [Vol. II,
coming from the ventral transverse connecting branch, the cau-
dal from the lateral tracheal tubes below the ganglion; one pair
supplying the middle part of the connectives between the suboes-
ophageal and the first thoracic; one pair supplying the first
thoracic and one unpaired to the same ganglion; one pair to each
of the remaining thoracic and one pair to each abdominal gang-
lion. The last ganglion receiving two branches from the caudal
region as well as two large branches from the system of the seventh.
The tracheae of the central nervous system of Corydalis larva
seem to be uniformly as outlined above, and the branches which
supply the nervous system come directly from large tubes and
these in most cases take their origin not far from spiracle openings
in the thoracic region and from very near the gills in the abdom1-
nal region.
The larger branches in the ganglia are quite uniform in the
distribution of their smaller tracheoles. There was found to be a
greater variation in the connective branches in the same individual
and in the ganglia of one specimen as compared with an other.
The supply to the connectives however, was quite uniformly given
by the tracheal systems of the ganglia at either end of the con-
nectives. The only variations were the constant ones; first, in the
additional extra supply to the long connectives between the sub-
oesophageal and the first thoracic ganglia; second, in the supply
to the connectives between the seventh and eighth abdominal
ganglia, where the tracheae from the seventh ganglion furnish the
connective branches.
All nerves connected with the ganglia were found to have a
good tracheal supply, usually one rather small tracheole which
branched many times and here as in the connectives, the branches
wound about without anastomosis and without any particular
indication of the direction of the nerve fibers, that is transverse
as well as longitudinal branches were numerous.
The branching of the smaller tracheae from the larger was rather
irregular and the course and distribution of the finest branches
was very complex. Within the ganglia as a rule fewer larger
branches were seen. On the surface, and each side of the middle
line dense masses of tracheoles outlined the central part of the
ganglion. But although there were these two dense tangles one
on each side in each ganglion, numerous minute, and some larger
branches crossed from one part of the ganglion to an other dis-
tant region and mingled with the branches from other systems.
1909] = Trachael Supply, Nervous System Corydalis ke
The small branches within the ganglion run straight or in broad
curves to a considerable extent, while the finer branches in the
connectives are often wavy. (Fig. M. & N.)
A few ends were found, but no clear case of anastomosis either
between systems or between branches of the same system,
although it looked like it in many places because of the great
length and intricate windings of the finest tracheoles. From the
smallest tracheoles also there were comparatively few branches
although single ones could be traced for long distances.
The connectives gave a good chance to study the distrihution
of the finer branches and their relation to each other. Here
although two systems overlapped, no anastomosis could be found.
(Fig. L.)
SUMMARY.
1. There is a very intricate and extensive supply of both
tracheal branches and capillaries to the central nervous system of
Corydalis larva.
2. There are three main branches to the brain on each side,
entering it laterally and ventrally.
3. There are two pairs of branches to the subesophageal.
4. There are three branches, two paired and one unpaired,
to the first thoracic ganglion.
5. There is but one pair of branches to all the other ganglia
except the eighth.
6. The eighth ganglion has two sources of tracheae. Two
branches from the seventh abdominal ganglion and two from the
caudo-lateral region of the animal.
7. The supply to the connectives in all cases but one, is from
the tracheal systems of the ganglia.
8. The supply to the long connectives between the subesoph-
ageal and first thoracic ganglion is in part from both of these
gangha and its middle portion is supplied by a small lateral tra-
cheal tube.
g. Nerves supplied by only one small tracheole may have
this forming a very complete network.
to. The frontal ganglion is supplied from the brain on each
side along the small connective and also from two small lateral
tracheae not connected with the brain.
tr. There is a superficial and a deep supply to each ganglion
and a very intricate network in each.
56 Annals Entomological Society of America [Vol. II,
12. In each ganglion, especially the abdominal, the central
area on each side is well marked by a dense tuft of minute trach-
eoles.
13. The large, and especially the small branches from one
system cross over to the regions of other systems to a marked
degree.
14. Although minute tracheoles are long and fine, no anasto-
mosis could be demonstrated.
15. Within certain lmits the tracheal supply is regular and ~
constant. ,
16. The tracheae supplying the ganglia are rather directly
connected with the larger tubes, not far from spiracles in the up-
per region and very close to gills in most of the abdominal region.
Comstock, J. H., and Kellogg, V. L., °95. The Elements of Insect Anatomy.
ithaca Ne we
Hammar, A. G., ’08. On the Nervous System of the Larva of Corydalis cor-
nuta L. Annals of the Ent. Soc. of America. Vol. I, No. 2, p. 105.
Krauss, W. C., ’84. On the Nervous System of the Head of the Larva of Cory-
dalis cornuta L. Psyche, P. 160. Pi. II.
I have especially to thank Prof. Kingsbury of the Department
of Histology, of Cornell University, and the Department of Ento-
mology of the same institution, for suggestions in the prepara-
tion of this paper.
From the Department of Histology and Embryology,
Cornell University, Ithaca, N. Y.
ANNALS E. S. A. Vor.. II, PLATE VIII.
W. A. Hilton.
ANNALS E. S. A.
Vor... II,
PLATE IX.
W. A. Hilton.
ANNALS, E. S. A. VoL. II, PLATE xX.
5 <i
IEE ee,
(ge 4 ee
je ae ae
ol >
W. A. Hilton
1909] Trachael Supply, Nervous System Corydalis 5
~I
EXPLANATION OF PLATES.
All figures indicated by letters are from photographs of the central
nervous system of Corydalis cornuta larva. The dark lines and darker parts
of the figures indicate tracheal tubes or dense masses of tracheoles.
Plate VIII.
Fig. A. Brain and part of the subesophageal ganglion from below.
xX 23.5.
Fig. B. Subesophageal ganglion from below. X 23.5.
Fig. C. Second thoracic ganglion from below. X 23.5.
Fig. D. Third thoracic (larger), and first abdominal ganglion from below.
Noa,
Fig. E. Abdominal ganglion from below. X 23.5.
Fig. F. Seventh and eighth abdominal ganglia from below. X 23.5.
Plate IX.
Fig. G. Two thirds of the brain from below. X 50.
Fig. H. Subespohageal ganglion from below. 50.
Fig. I and J. Abdominal ganglia from below. 50.
Plate X.
Fig. K. Upper portion of a thoracic ganglion showing larger branches only.
x 50
Fig. L. Part of two connectives showing the mingling of two tracheal systems
ineach. X 80.
Fig. M. Abdominal ganglion from below at one focus. > 80.
Fig. N. Abdominal ganglion from below, same specimen as Fig. M, only ata
different focus. > 80.
PLATES XI AND XII.
Camera lucida drawings of the main tracheal branches in each of the ganglia
of larval corydalis. Chiefly from a single specimen. Each is drawn from below
and the scale at the left is equal to one half mm.
Plate XI.
Fig. 1. One half of the brain showing the main branches.
Fig. 2. One half of the brain of another specimen, showing most of the branches
that can be seen from one side.
Fig. 3. Subesophageal ganglion.
Fig. 4. First thoracic ganglion.
Fig. 4. Second thoracic ganglion.
Fig. 6. Third thoracic ganglion.
Plate XII.
Fig. 7. First abdominal ganglion.
Fig. 8. Second abdominal.
Fig. 9. Third abdominal.
Fig. 10. Fourth abdominal.
Fig. 11. Fifth abdominal.
Fig. 12. Sixth abdominal.
Fig. 13. Seventh and eighth abdominal ganglia.
Vor. il) Bran 20K.
ANNALS E. S. A.
W. A. Hilton.
EVOLUTION AND ADAPTATION IN THE PALPUS OF MALE
SPIDERS.
Jas. A. NELSON.
The following conclusions are the results of study of 85 repre-
sentatives of 21 families of American spiders, taken from the col-
lection in Cornell University, through the courtesy of Professor
Comstock.
The simplest form—structurally—of the male palpus is that
found in the sub-order Tetrapneumones, which, as is well known,
is recognized as including those members of the Araneida which
are primative in respect to other structural characters. It will
therefore be justifiable on two grounds, namely, the structure of
the palpus itself, and the general structure of the members of this
group—to regard this form of male palpus as the most primitive
existing type, thus enabling us to form some conception of its
ancient condition. It accordingly serves to indicate the point of
departure from which the other more highly modified types have
beendeveloped. This simpler type of male palpus 1s fairly well de-
fined, although varying in many particulars in the different mem-
bers of the sub-order. Fig. 1 isa schematized representation of this
type. The essential portion of the palpal organ, the bulb (6), is
placed at the end or on the ventral surface of the sixth, tarsal, or
terminal segment, if we consider the female palpus as represent-
ing the original form of the second pair of appendages. The bulb
varies from long pyriform to spheroidal in shape; it is articulated
at its base to the tarsal segment by a narrow zone of unchitinized
integument, and bears at its summit a tapering process, or style
(s). Within the bulb is a tubular canal, coiled in a helicoid curve.
This canal is wider at its base and narrows gradually toward its
peripheral end, which passes into the style and opens at its apex
by a narrow aperture. Its function is to contain the sperm and
it is therefore termed the semeniferous canal (represented in
dotted outline). It is important to note the position of the heli-
coid curve just mentioned. It is placed in such a manner that
its central axis, the helical line (represented in the figures by an
arrow), passes through the base of the style at the distal end of
the bulb, and the base of the bulb itself at its central end, in this
60
1909 | Evolution in Palpus of Spiders 61
primitive type, and moreover it generally coincides with the
long axis of the bulb, whenever this is in evidence. This line or
axis is a valuable basis for orientation in the manifold changes in
form undergone by the bulb in the higher families, and I will,
for convenience, term it the princtpal axis of the bulb. This axis,
as the figure shows, generally forms an angle of from 50 to 90
degrees with the long axis of the tarsal joint. Four characters are
to be noted as marking this type of palpus: (a) the more or less
cylindrical form of the tarsal joint; (b) the position of the bulb,
frequently terminal and rarely, if ever, near the base of the tarsus;
(c) the narrowness of the zone of articular membrane joining cym-
bium and bulb, scarcely wider than that between the joints of
other appendages; (d) the integument of the bulb itself, which is
a continuous chitinized sheet, not broken up into segments or
plates. This type of bulb I propose to designate by the term
holochitinous.
Turning to the palpal organ of a representative of the higher
spider families in the Dipneumones (figs. 2-7) it is seen that the
primitive type just described has undergone great alteration,
especially in regard to one or more of the four characters des-
cribed. These are, broadly speaking, the following: (a) change
in the form of the tarsal segment or cymbium; (c) this has be-
come compressed in a plane at right angles to that in which the
bulb lies, so as to present an elliptical or oval outline in face view,
and is moreover hollowed out beneath the base of the bulb to
form the alveolus (a); (b) The position of the bulb is uniformly
near the base of the cymbium, never, in the higher groups, near its
apex; (c) the articular membrane joining bulb and cymbium
has increased greatly in extent, and has become an elastic sac
(figs. 6 and 7) capable of being greatly distended by blood pres-
sure, thus erecting the bulb, thrusting it out some distance from
the alveolus and away from the cymbium. It is accordingly
termed the haematodocha; (d) The bulb itself has suffered alter-
ation in that it is now generally split transversely into two or
more segments, united by membranous integument. It is also
frequently greatly shortened in its principal axis.
Each one of the above mentioned alterations of the primitive
type seem quite evidently directed toward one end, namely,
protection of the bulb, the essential part of the copulatory organ,
from mechanical injury. Taking the above four named altera-
tions in order: (a) Change in the shape of the tarsal joint, from
62 Annals Entomological Society of America [Vol. IT,
cylindrical to a flat boat-shaped form, hollowed out on the side
toward the palpus, would very evidently be of service in shield-
ing the bulb from mechanical or other injuries.
(b) Change in the position of the bulb, from an exposed posi-
tion at or near the tip of the cymbium, is correlated with the
change in the form of the cymbium mentioned under (a), and
enables the bulb to make full use of this alteration.
(c) The distensibility of the articular membrane makes a long
or projecting bulb less necessary, since it may be projected at the
time of copulation, and only then.
(d) The purpose of splitting the bulb into segments or plates
is, in the majority of cases, less evident, but in one large family,
the Lycosidae, it enables the bulb to be closely folded together
in the alveolus.
The whole evolution of the male palpus, therefore, appears
to hinge on the one adaptation, and the different modifications
in the form of the palpal organ are accordingly only illustrations
of the different means which may be employed to serve this end.
Above all stands out the constant mechanical adaptation or fit-
ting together of bulb and cymbium to one another, often carried
to an extraordinary degree. When the palpus is at rest, the bulb
and cymbium together, in the majority of the Dipneumones, form
acompact, more or less ovoid mass. This is brought about, gen-
generally speaking, in one of five ways, thus dividing the American
spider families of the sub-order Dipneumones into at least five
divisions. The lines of separation between these divisions, are,
of course, not always sharply marked, since transitional forms
occur, but the types themselves are nevertheless quite distin-
guishable. They are as follows:
1. First of all are those families in which the structure of the
male palpus differs little, if any, from that described for the
Tetrapneumones. These families are:
Filistatidae.
Dysderidae.
Sicariidae (Scytodes).
The Pholcidae are probably also to be placed here. It is to
be noted, that the cymbium of Dysdera crocata shows evidence
of the flattening so constantly present in the higher families.
2. Fig. 2. The principal axis is greatly shortened, the bulb
is in cross section nearly circular, and sinks into the circular
1909| Evolution in Palpus of Spiders 63
alveolus like a ball into a socket. The principal axis is approxi-
mately perpendicular to the long axis of the cymbium. Repre-
sentatives:
Thomisidae (in part).
Agalenidae.
Pisauridae.
Oxyopidae.
Dictynidae.
3. Fig. 3. This is apparently derived from the second type.
The principal axis is not only much shortened, but the bulb is
elongated in a new axis at right angles to the principal axis and
parallel to the long axis of the cymbium, the alveolus being cor-
respondingly extended so as to receive the basal half of the bulb,
and therefore oval or elliptical in outline. Representatives:
Thomisidae (in part).
Clubionidae.
Gnaphosidae.
4. Figs. 4, 6, and 7. The alveolus is extended so as to
include practically the entire ventral face of the cymbium, reduc-
ing the latter to a mere concave shell. In the closed palpus the
distal portion of the cymbium lies against the side of the bulb, in
other words, the principal axis forms an angle much less than go
degrees with the long axis of the cymbium. Representatives:
Therididae.
Argiopidae.
Mimetidae.
Uloboridae.
5. Fig. 5. The chitinized covering of the bulb, particularly
of its distal end, is reduced in extent, so that the bulb is largely
composed of thin membrane and is therefore collapsible, in its
inactive condition being folded up into a flat package, almost
completely contained in the shallow alveolus, which, as in the
first division, is moderate in size. The principal axis is slightly
oblique to the long axis of the cymbium. Represented only
by the
Lycosidae.
These modifications in the form of the bulb and cymbium,
which appear so clearly to be designed for the protection of the
former, are aided by special modifications or structures directed
64 Annals Entomological Society of America [Vol. II,
toward the same end. Two of these are worthy of mention.
They are: The accessory branch of the cymbium found in the
Linyphiinae and Tetragnathinae, sub-families of the family
Argiopidae, (figs. 6 a. b.), and the expanded patella of the Theri-
diidae (fig. 7 p). Both of these structures serve to overlie and
protect the base of the bulb.
To conclude: The more complex palpi found in the Dip-
neumones have been developed by modification of a palpus
similar to that of fig. r. and the principal factor in their evolution
has apparently been that of protection from mechanical injury.
EXPLANATION OF PLATE XIII.
Fig. 1. Diagram of o palpus of the most simplified form.
Fig. 2. o& palpus of Misumena, slightly digrammatic.
Fig. 3. o palpus of Phurolithus, from camera drawings.
Fig. 4. o palpus of Araneus, schematic.
Fig. 5. < palpus of Lycosa, also schematised.
Fig. 6. oo palpus of Erigone autumnalis, from a camera sketch.
Fig. 7. G palpus of Lithyphantes corollatus, also from a camera sketch.
The strongly chitinized portions of the palpus are represented in heavy
outline, the membraneous portions, including the haematodocha, in lighter out-
line. The semeniferous canal is represented by dotted outlines, the principal
axis, in each case, by an arrow. The morphological ventral side is turned up-
ward.
a, alveolus; a. b., accessory_branch; b, bulb; c, cymbium; /#, haematodocha;
p, patella.
Volume II. Number 2.
ANNALS
OF
The Entomological Society of America
JUNE, 1909
EDITORIAL BOARD
J. H. COMSTOCK, L. O. HOWARD,
Irwaca, N. Y. WASHINGTON, D. C.
Coa oe BE DEUINE, W. M. WHEELER,
GUELPH, ONTARIO, CANADA. Boston, MAss,
C. W. JOHNSON, P. P. CALVERT,
Boston, Mass, PHILADELPHIA, Pa.
V. L. KELLOGG, J. W. FOLSOM,
STANFORD UNIv., CAL, URBANA, ILIS.
HERBERT OSBORN, Managing Editor,
COLUMBUS, OHIO.
PUBLISHED QUARTERLY BY THE SOCIETY
Entered as second class matter April 11, 1908, at the Post Office at Columbus, Ohio,
under the Act of Congress of March 3, 1879.
Publications of the Entomological Society of America.
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Proceedings of first three meetings; Constitution, By-Laws and List of
Mem bersa eee rinants pte reinve ce tecarekeDledevertie tacks coe ehclad heenabetettepeteteter mevctan te bstevenh .25
WHEELER, Wa. M-—Polymorphigm |/on (ANS...) cei chelelajeieyeleiste viele telels/el setalnes .30
OsporNn, Herbert—The Habits of Insects as a Factor in Classification...... .20
Severin, H. H. anp Sreverrn, H. C.—Anatomical and Histological Studies
of the Female Reproductive Organs of the American-Saw fly, Cimbex
A TNENICAN ALGAE 4 ately «des ata e ele Vpeletey eis stolona te bale ohielealaia fe tn areas ue Peat .25
Feit, E. P.—Some Problems in Nomenclature...............ceeesecceeess 10
Hammar, A.G.—On the Nervous System of the Larva of Corydalis cornuta L .25
Brapiey, J. C.—A Case of Gregarious Sleeping Habits among Aculeate
IEP VINENO PLera ANNs ae ese (ok Wiel ated LUM eC Lelstettelel ANAL MaiL ee ite wie Ma Re aan telat -10
Davis, J. J—Notes on the Life History of the Leafy Dimorph of the Box-
elder Aphid, Chaitophorus negundinis Thos................eeeeeeeeee -10
HamBuetron, J. C.—The Genus Corizus, with a Review of the North and
MiddlevAmericunsS peace: 2 rack MV leverotbnisiataie/eters ele ciavesce Sicha s elolers -25
Girau.t, A. A.—Biological Notes on Colorado Potato Beetle............... .25
Girautt, A. A.—A Monographic Catalogue of the Mymarid Genus Alaptus .25
Severin, H. H. anp Severin, H. C.—Internal Organs of Reproduction of
Male Saw-tlye in ie ran deve oe tie aie vont ca averat ta aetna ae woke (avaya/ a cM Cee Me pec ave ts 15
Smitu, C. P.—A Preliminary Study of the Aranez Theraphosae of California .75
DAVIS, J... Studien On WApMIGidae 2) lily eile siete cee tatnraysiadentioeinin es sete .20
Ritzy, W. A.—Muscle Attachment of Insects. .............0....ceeccceees 15
Nerpuam, J. G.—Critical Notes on the Classification of the Corduliinae
COCO ata) Hae ee Ee MI ate le Ste LoaL Rt otLkc, i tage Date is REMRM G RTL iota Mae ane ale eh LO
Howarp, L. O.—A Key to the Species of Prospaltella with Table of Hosts
and Descriptions of Four New Species............cceeecseccccsecceces 15
Hoop, J. D—Two New Species of Idolothrips.............. 2... .eeeee eens -10
Address
ANNALS ENTOMOLOGICAL SOCIETY OF AMERICA,
Biological Building, O. S. U., Columbus, Ohio.
ANNALS
©}F
The Entomological Society of America
Volume II PUNE; 1909 Number 2
INVESTIGATIONS OF TOXOPTERA GRAMINUM AND ITS
PARASITES.
By F. M. WEBSTER,
BUREAU OF ENTOMOLOGY, WASHINGTON, D. C.
This minute insect (see figs. 1, 2), aside from its economic
importance, which, by the way, is very great, has become the
basis of a long series of thorough investigations, carried out by
several assistants of the Bureau of Entomology, under my direc-
tion. My own studies of the species were made largely during
the years 1884 to 1890, while employed by the old Division of
Entomology. As the subject of this paper may not be familiar
to all entomologists it will be better to begin, historically at least,
with our first knowledge of the species in its native country,
if, indeed, we might with our present information, be allowed to
designate for it a place of nativity.
__ Fic, 1.—The spring grain-aphis (Toxoptera graminum): a, Winged migrant
viviparous female; 6, antenna of same. a, Much enlarged; b, highly magnified.
(From Pergande. )
67
68 Annals Entomological Society of America [Voli
OCCURRENCE IN THE EASTERN HEMISPHERE.
It seems to have been observed at Parma, Italy, as early as
1847 (1). Five years later, in 1852, (1) Rondani, who described
the species during this year, wrote to Professor Bertoloni under
date of June 14, also from Parma, relative to the insect as follows:
“We have in our city an innumerable number of insects of a
species of the Aphis genus, of Linnzeus, of the order of Hemiptera.
‘
Fic. 2—The spring grain-aphis or ‘“‘green-bug’’ (Toxoptera graminum: a,
wingless female; b, larva;c, pupa. Much enlarged (from Pergande.)
Sometimes and in certain places the number of these insects fly-
ing in clouds in the air has been so great as to render them troub-
lesome to people, entering the nose, eyes, and even the mouth
when one cannot think how to protect oneself from them.”
Elsewhere in this letter Rondani states that he had never been
able to find it on any but graminaceous plants, where it nestled
on the leaves. In commenting on this letter of Rondani, (1)
Prof. Bertoloni takes occasion to say that:
‘“Innumerable specimens of the Aphis graminum Rondani,
are seen in the streets of the city of Bologna and these have sev-
eral times entered my nose and eyes when passing rapidly along
the canal of Reno.”
Passerini (2) excepted we find no mention of the insect
again until 1884, when Dr. G. Horvath (3) mentions an attack
on oats in Central Hungary in June, 1883. ‘Ten years later, in
1894, Professor Karl Sajo, (4) records another outbreak, also in
Hungary, again among growing oats.
Schouteden (5) in 1906, records the species from Belgium,
but gives no further data except that it affects the graminacee.
1909] Toxoptera graminum and its Parasites 69
In August, 1907, at the meeting of the International Society of
Zoologists in Boston, and, later, also in Washington, I had the
good fortune to meet Dr. Horvath and from him learned that the
species had been found in Siberia on graminacee.
For two or three years past, grain in the Orange River Colony,
South Africa, has been seriously affected by a species of Aphis,
we having received several complaints of these invasions from
Mr. H. Neethling, at that time, Chief of the Horticultural and
Biological Division of the Department of Agriculture at Bloem-
fontein. It was only within the last few months, however, that
we were able to secure specimens of these aphids for identification
and, surprisingly enough, they prove to be Toxoptera graminum.
With these were a number of individuals of a species of Lyszphle-
bus, allied to the one that is so efficient in holding the pest
in check in America. This gives us records of the occurrence of
the species in Europe, Asia and Africa.
OCCURRENCE IN THE WESTERN HEMISPHERE.
The earliest information we have of the occurrence of Toxop-
tera graminum in America is in the year 1882, when it was received
at the Department of Agriculture, but sad to say there is no
record as to where the specimens came from. Late in June, 1884,
it was found sparingly on wheat in the vicinity of Cabin John,
a short distance north of Washington, and, about the same time,
I found it infesting wheat plants in some breeding cages where
some experiments were being carried on by myself at Oxford,
Indiana, about one hundred miles southeast of Chicago, Illinois.
Nothing more was heard of the species until 1890 when a very
serious and destructive outbreak occurred in the West, extend-
ing from northern Texas to Missouri and central Indiana. Cur-
iously enough there was little complaint from the territory east
of the Appalachian mountain system, even where considerable
wheat and oats were being at that time grown. Again, we have
a period of eleven years without noticeable injuries from this pest;
then, in 1901, there came the most destructive outbreak that had
occurred up to that time, but in this instance depredations did
not extend farther north than central Oklahoma, though planters
in South Carolina afterwards reported ‘‘green lice’? as having
been very abundant on oats during the spring of that year.
In March, 1903, there was an incipient outbreak of the species
in northern Texas. But this seems not to have extended north-
ward beyond the Red River and was overcome by parasites in
70 Annals Entomological Society of America [Volmit:
April. (6) This time it certainly did appear in excessive abund-
ance on fall-sown oats in North and South Carolina, and planters
became alarmed over its presence. It did not, however, work
serious injury there.
In December, 1g06, the insect began to appear in fields of fall
sown and volunteer oats in northeastern Texas. Throughout the
winter of 1906-1907 they increased in abundance and by spring
had ravaged the fields of both wheat and oats throughout Texas
and Oklahoma, east of the 100° meridian, and as far north as
Wichita, Kansas, but spring sown oats were damaged as far north
as eastern Iowa, northern Illinois and:southern Wisconsin. In
the east there was also serious damage to wheat and oats in both
of the Carolinas. The loss by reason of this outbreak would
probably not fall short of $12,000,000 to $15,000,000 and proved
to be much more serious than any preceding occurrence. Thus
it will be observed that the species is becoming constantly more
abundant, its re-occurrences more frequent and more threatening
to grain crops, especially throughout the country between the
Rio Grande and Missouri rivers. ‘This is also true to a less degree
in the Carolinas.
DISTRIBUTION IN AMERICA.
Most rigid searches have failed to reveal the presence of the
insect in the country between the St. Lawrence river, Lake Onta-
rio and Lake Erie, and the coast, to a line drawn from Cleveland,
Ohio, to Philadelphia, Pa., although it would not be surprising
if it extended farther northeast nearer the sea coast.* Neither
has it been found in northern Michigan, northern Wisconsin,
northeastern Minnesota, or in the arid region of western North
and South Dakota or eastern Montana, where it has also been
diligently searched for. It has not been found in Alabama, Miss-
issippi, or Florida, but has not been searched for in those States.
With these exceptions it ravages over the whole country east of
the Rocky Mountains, from Mexico northward into Manitoba
and Saskatchewan, and from an elevation little above sea level,
to 8,o00 feet in New Mexico. On the Pacific slope it is found in
astern Washington. Whether it also extends from Washington
southward along the Pacific coast to Mexico is not known,
because it has not been searched for in that portion of the
country. (See map.)
* Mr. Paul Hayhurst, assistant to Dr. Wheeler, states that he has found it
on oats, Agropyron repens and Dactylis glomerata, in the vicinity of Forest Hills,
near Boston, Mass.
1909] Toxoptera graminum and its Parasites a
If we assume that Southern Europe is the original home of the
Species, its occurrence in the western hemisphere can only be
accounted for on the score of introduction, but the distribution
of the species in the eastern United States, does not in the least
indicate such an introduction as having come through any of the
Atlantic coast seaports. On the other hand, its occurrence all
along the Rio Grande River bordering Mexico, together with its
destructive abundance in northern Texas, as early as 1890, only
Z (oy
ST \Y
f *
LISTRIBUT/ON OF TOXOPTE RA GRAMINUM IN THE UNITED STATES.
eight years after its first discovery in America, goes far to streng-
then the theory of an early introduction into some country to the
South, and a natural diffusion over the United States, following
the same lines of dispersion as for so many other species coming
to us from Mexico and southward. .
FOOD PLANT.
The species is essentially a leaf infesting insect, rarely infest-
ing the stem. While preferring oats it will readily attack wheat,
rye, barley, and may often be found on the underside of the lower
leaves of corn. Corn excepted, its effect on the foliage of grain,
when present in large numbers, is to cause the infested leaves to
change to a red color, which seems to be very characteristic of
Toxoptera and does not follow, to a marked degree, attacks of
72 Annals Entomological Society of America [Vol. II,
other species of Aphids on these grains. The insect also has
been found breeding upon a considerable number of native grasses
any one of which may constitute its alternating food plant, upon
which it may survive the summer in different portions of the
United States. It has been found breeding freely upon Alope-
curus geniculatus in Oklahoma by Mr. W. J. Phillips and by Mr.
C. N. Ainslie in Kansas; on Agropyron occidentalis, also in Okla-
homa, by Mr. Phillips; and by Mr. O. E. G. Kelly and Professor
Gillette in Colorado. Agropyron tenerum was found moderately
infested by Mr. C. N. Ainslie at Las Vegas, New Mexico. It was
found breeding upon Bromus at Washington, D. C., and also
upon Bromus portert at Las Vegas, New Mexico, and on an un-
determined species of Bromus at Mesilla Park, New Mexico, also
by Mr. Ainshe. I observed it very abundantly on Dactylis glom-
erata in Indiana in 1890, and again excessively abundant in a
small, isolated meadow, of this grass near Midlothian, Virginia,
in April, 1907. This meadow was located in a region not adapted
to the growing of grain and there was no field of growing wheat
or oats within five miles. Mr. Kelly found it inhabitating Dzs-
tichlis spicata in such abundance as to be damaging this grass in
Montana, which in that part of the country is known commonly
as ‘‘salt grass.”’ It was found inhabiting Elymus striatus at Las
Vegas, New Mexico by Mr. C. N. Ainslie. Mr. W. J. Phillips
found it attacking Hordeum pusillum at Beloit, Kansas, and,
Kingfisher, Oklahoma, while I found this to be of frequent occur-
rence throughout Kansas. Mr. Kelly observed it abundant on
Hordeum jubatum in Montana, while Mr. C. N. Ainslie found it
moderately abundant on Hordeum cespitosum near Cimarron,
New Mexico. Wherever Poa pratensis grows the insect will prob-
ably be found breeding upon it through the entire summer. In
fact this has been actually observed to occur in the city of
Washington. Mr. Phillips has also observed it in Indiana and
throughout the northern sections of the country, where Toxoptera
occurs; it is more likely to be found on this plant during summer
than upon grain, excepting, perhaps, during seasons of excessive
abundance. Mr. C. N. Ainslie also found it moderately abundant
on Polypogon monspeliensis about Albuquerque, New Mexico.
It has been found breeding on Stipa viridula at Las Vegas, New
Mexico, by Mr. C. N. Ainslie, the grass being heavily infested.
As one or more of these grasses will be found to occur in almost
every portion of the United States, it would appear that the non-
1909] Toxoptera graminum and tts Parasites 73
occurrence of Toxoptera in any considerable section of the country
cannot be due to a lack of uncultivated food plants. Mr. Phil-
lips found that the species readily developed on the following
plants in his breeding cages: Dactylis glomerata, Eleusine indica,
Eragrostis pilosa, E. megastachya, Sporobolus neglectus, A gropyron
sps., Elymus virginicus, E. canadensts and bromus secalinus.
SEASONAL HISTORY.
Though in the main following quite closely the developmental
habits of other Aphids, this species exhibits some striking con-
trasts. While it probably passes the winter in the egg state in
the northern portions of the country, it is quite certain that it is
not restricted to that mode of hibernation during mild winters,
or in the South. The fact that viviparous 92 9 (figs. 1, 2)
sent from Leavenworth, Kansas to Washington, D. C., in March,
and there placed indoors, produced oviparous 2 2 (fig. 3) and
oo (fig. 4) in April, and, in case of the latter, even till May 18,
shows that the sexes may occur in spring. As information on
the occurrence of the sexual forms appears to be entirely lacking
in the South, it would be too much to connect the presence of
these forms in spring as showing a changed summer condition
in the South, whereby the dry season instead of the winter
might be passed in the egg. Bearing upon this point, but prov-
ing nothing unfortunately, it may be stated that at Richmond,
Indiana, in the autumn of 1907, Mr. Phillips encountered a
pronounced lack of oviparous 2 2, the weather being unusually
wet. During a corresponding period of 1908, with one of the
severest droughts on record prevailing at the time, he was unable
to get a sufficient number of viviparous
2 2 to. keep up his experiments, while
the sexual forms were present in abun-
dance.
Two of my assistants, Mr. W. J.
Phillips, and Mr. C. N. Ainslie, have on
occasions dissected 2 2 and found both
eggs and embryos in the ovaries, the
latter in spring; while the former, in
the fall, observed in one instance that
; : Fic, 3—Spring grain-aphis
the eggs were in one side and the em- (Toxoptera ete On
bryos in the other side of the abdomen, pPatous female with eggs in
body, greatly enlarged; at
as though one ovary had _ produced right? egg stili more enlarged.
eges and the other embryos. (Original. )
vA Annals Entomological Society of America [Vol. IT,
Mr. Phillips in a long series of breeding experiments carried
out at Richmond, Indiana, has found that the oviparous 2
will not deposit eggs without having first mated with the oJ,
and usually produces not more than five or six eggs. The
life of the viviparous 2 is much longer than that of the oviparous
2 but the life of the latter is greatly prolonged if she fails to
meet the co’ and become fertilized.
HIBERNATION.
The species will breed at about or a little above 45° Fah.,
though, as determined by Mr. Phillips, they do not reach their
maximum until the temperature rises to above 50° Fah., which,
however, is still too low to render their parasites active. As an
experiment, a miscellaneous lot of individuals were frozen up in ice
Fic. 4—Spring grain-aphis (Toxoplera graminum: Male.
Greatly enlarged. (Original.)
for five, twelve and twenty-four hours respectively, and on being
thawed out the older individuals all succumbed; in another twen-
ty-four hours experiment, using individuals of different stages
of development, only the pupz of the viviparous 92 survived. |
Viviparous 2 2 brought indoors from a temperature of 11° Fah.,
and thawed out began to give birth to young in about three hours.
Thus it is that the species is able to withstand the climate of
northern latitudes and high altitudes. As it breeds freely in a
temperature of roo° Fah., in the shade, presumably still higher
in the open, on the public grounds in Washington, it would seem
that farther south, where it was found by Mr. C. N.-Ainslie, the
insect would encounter even a greater degree of heat in summer.
Thus we have a fairly good basis for the supposition that it
1909] Toxoptera graminum and its Parasites 75
extends southward beyond the boundaries of the United States.
90 far we have not found the sexual forms south of southern Kan-
sas, about latitude 37°, but north of this latitude it can certainly
pass the winter ordinarily in the egg state and in mild winters as
viviparous 2 2 either wholly or in part developed. South of
Lat. 37° there really seems to be no vital necessity for sexual 2 Q
in order to perpetuate the species or to enable it to pass the winter.
Indeed, the greatest need would seem to be for some means of
bridging over the dry period, during which there is a serious lack
of food and in a temperature far too high to induce torpidity thus
rendering food unnecessary.
DEVELOPMENT DURING SUMMER.
Beginning July 8, 1907, Mr. W. J. Phillips, at Richmond, Ind.,
began a study of the continuous generation of this species, follow-
ing it through to December ro, besides securing sexual 2 2 and
eggs therefrom on blue-grass in the fields in October. With these
eggs, he began March 27, 1908, to again follow out continuous
generations, commencing with the first stem mother that hatched
from eggs deposited the previous autumn, From the wintered
over egg to the oviparous 2, or from one egg stage to the other,
the maximum was twenty-one generations of viviparous 2 2
when the first born were used in each case, but only ten genera-
tions where the latest born was used in a parallel experiment.
The experiment ended December 1, 1908. It is not unlikely that
there are a greater number of generations produced in the south,
where the warm period is more prolonged, though in the north
this period while shorter is not usually so excessively dry.
Ordinarily, the viviparous 2 will reach full development and
herself begin reproduction about eight days from birth, but in late
autumn this period becomes very irregular. In one case Mr.
Phillips had a viviparous 2 under observation, under outside
conditions from the day of her birth October 18 to December ro.
Though apparently fully developed she produced no young and
went into winter in a most vigorous condition. It will be noted
that it was just such as these that survived the freezing experiment
previously described. The young, when born, are enveloped in
a membraneous sack, from which they disengage themselves
almost the instant they leave the mother. A viviparous 2 may
produce as many as 70 young scattering them throughout a period
76 Annals Entomological Society of America [Wolk via:
of four to six weeks. The wingless viviparous 2 2 are more pro-
lific than the winged, but, of course, scatter their young less as they
are confined to a very limited area. It is usual to observe a wing-
less mother stationed on a leaf in the midst of a family of a dozen
or more young, varying in size from newly born to half grown.
The insect moults four times.
EFFECT OF METEOROLOGICAL CONDITIONS.
As has already been shown, the direct effect of low temperature
on the insect is, in the south at least, exceedingly small. It is
vastly more probable that the influence of winds and humidity
are much more important factors, though of the latter we cannot
yet speak with the fullest assurance.
As has also been shown, it may turn out that hot dry weather
may be found to greatly influence the time of appearance and
abundance of the & and sexual 9.
Winds certainly do have considerable influence in the disper-
sion of the winged viviparous 2 &, during the seasons of excessive
abundance, and as it is well known that a diminution of the food
supply, lke the killing of the grain plants, tends to produce
winged instead of wingless individuals, it will be seen at once that
the occurrence of these winds at the time of a serious outbreak
may become a powerful factor in the spread of the pest. Now,
the prevailing winds in the Southwest during spring are from a
southerly direction and as there is a continually increasing food
supply to the northward as the season advances, these winds help
the insects to keep pace with this advance. This diffusion might
be likened to a horizontal revolving movement, the winged 2 92
from a destroyed field passing over others along the border and
settling down to breed, infest and destroy other fields more in
advance, and so on day after day, thus passing over miles upon
miles of young growing grain changing it from a luxuriant green
to reddish brown, as effectually destroying it as would be the case
were it to be scorched by fire. There are of course other modes
of dispersion, and I have observed not only winged individuals
but also parasites, clinging to the glass of the windows in railway
coaches and in this way carried long distances.
Thus it is that an invasion of Toxoptera, originating in central
northern Texas in a winter temperature above the normal, yet
too cool to admit of its chief natural enemy becoming active, will
1909] Toxoptera graminum and tts Parasites aT
develop countless millions of winged viviparous 2 2, and these
will gradually become diffused northward as the season advances
and weather conditions favor them. Indeed the experience of
Rondani and Bertoloni, in Italy, found an equal in the southwest
in 1907 where Mr, C. N. Ainslie observed the winged 2 2 in such
swarms as to interfere not only with a Sunday game of baseball,
but also with the conducting of a funeral. North of about Lat.
38° the season is so far advanced that wheat becomes too large
and tough to offer the requisite food supply, but spring oats here
prove a convenient and attractive supplement and it is usually
this crop that suffers most to the northward, provided, of course,
the weather conditions are not favorable to the development of
natural enemies.
NATURAL ENEMIES.
There are a number of these, especially among the Coccinelli-
dz, several species of which feed, both in the larval and adult
stages, on Toxoptera. Syrphus flies and Chrysopa also destroy
them. A phelinus nigritus Howard, recently described, is a min-
ute parasite that has been reared from this insect 1n South Caro-
lina by Geo. G. Ainslie, and in New Mexico by Chas. N. Ainslie,
both assistants in the Bureau of Entomology. Quails are very
fond of them and Miss Margaret Morse of Clarke University has
been kind enough to make some experiments for us in feeding these
birds. Miss Morse estimates that about 5,000 individual Toxop-
tera were eaten by a single quail in one day, preference being
shown for those that were unparasitized. The Song Sparrow,
Melospiza melodia, also devours great numbers of them in the
grain fields.
DEVELOPMENT AND INFLUENCE OF LYSIPHLEBUS TRITICI.
As a matter of fact, however, all of the natural enemies pre-
viously mentioned are of small moment as compared to the influ-
ence of the one minute parasitic species Lystphlebus tritici (figs.
5, 6.). It is this species, or what we are at present terming as
such, that normally holds Toxoptera in check in this country, and
so long as its development and activity are not obstructed by
meteorological conditions, it will probably continue to do so.
Indeed so important is this insect and so powerful is its influence
that only a short space of from ten days to two weeks time 1s
78 Annals Entomological Society of America [Vol. II,
required for it to overcome a most serious outbreak of Toxoptera
and thus save from destruction vast areas of growing grain. The
species winters over in
the fields in the bodies
of its host. In many
cases this hibernation
is passed as nearly or
quite fully developed
adults, ready to
emerge when the tem- Fic. 5.—-Lysiphlebus depositing its eggs in the
perature rises to the body of a grain-aphis. Much enlarged (original.)
neighborhood of 56° and continues for a sufficient length of time,
and where they have been prevented from emerging the pre-
vious fall, on account of the advent of cold weather. This is clearly
shown by the fact that Mr. E. O. G. Kelly found them in this
condition at Leavenworth, Kansas, on November 13. From a
lot of 50 parasitized Toxoptera that had been washed or rubbed off
Fic. 6.—Lysiphlebus tritict, principal parasite of the spring grain-aphis;
Adult female and antenna of male. Greatly enlarged (original). At right: Egg
of Lystphlebus tritict. Highly magnified (original).
the leaves of the young grain (fig. 7) and were taken out of the
mud about the wheat plants on February 28, after the winter was
practically over, Mr. Kelly found that of these 50 dead bodies,
17 contained full grown larve of the parasite, 12 contained pupae
of a light color and 21 contained pupz of a dark color, the latter
T909| Toxoptera graminum and its Parasites 70
evidently ready to promptly develop with the advent of warm
weather. Mr. Kelly, on the same date, also secured a large num-
ber of Toxoptera in various states of development that were hiber-
nating in wheat fields near Leavenworth, Kansas. The weather
had been such as to preclude the possibility of these having been
recently parasitized. Yet some of them soon began to show the
characteristic yellow color of a Toxoptera parasitized by Lysiphle-
bus, and adults were afterwards reared from them. This shows
conclusively that Lystphlebus hibernates in advance stages of
development in the bodies of its host, which they have killed the
previous autumn, as well as larve in hosts wintering over from
half to fully grown. |
Fic, 7.—Stalk of wheat, the leaves covered with dead spring grain-aphids
killed by the parasite Lystphlebus tritict. About natural size (original).
80 Annals Entomological: Society of America [Vol 1g
The 2 Lystphlebus is even more prolific than Toxoptera. Mr.
Phillips has found 2 2 with upwards of 4oo eggs in their ovaries,
and Mr. Kelly has reared in some cases 206 individuals from a
single mother Lystphlebus. The eggs are lemon shaped and white
in color. When excessively abundant the parasite will thrust its
ovipositor into old and young aphids of both sexes, including the
sexual 2, even though previously parasitized; and Mr. Phillips
has observed them to apparently oviposit in the dead bodies of
those that had been killed by fungus attack. When their num-
bers are not so great, they show more discrimination and seem
to prefer half grown individuals in whose bodies to place their
eggs.
This species of Lystphlebus is parthenogenetic, as was first
observed by Mr. Phillips at Richmond, Indiana, and afterwards
more fully elucidated by Messrs. Kelly and Urbahns at Welling-
ton, Kansas. The experimental breedings of Mr. Phillips in 1907
indicated that the offspring of virgin 2 Lystphlebus were always
exclusively co'c. In a series of upwards of eighty breeding
experiments carried on indoors, in 1908, by Kelly and Urbahns,
only forty-eight gave results of any kind. Of these only four
gave 2 9, the other giving only oc’. In the four exceptional
cases the 2 @ kept virgin, ran all to oo’, two with the first
generation, one with the second and one with the third. The
mode of procedure was as follows:
Starting with a mated 2, the 2 2 from among ae offspring
were isolated, even before emergence; on their appearance these
were given Toxoptera reared under cover to preclude parasitism;
the few 2 2 from among this second generation were again iso-
lated in the same manner, the 2 @ in all cases being kept un-
mated. It was thus found possible to breed a limited number
of 2 2 parthenogenetically to and including the third generation.
Beyond this all offspring were o'<c, this seemingly being the
limit. Just why such a large percentage of these experiments
should have proven abortive is not clear. The conditions under
which they were carried out were of course unnatural but much
more protected from the adverse elements of the open field.
The egg of the Lystphlebus normally develops to the emerging
adult in about ten days, during the first six of which the host
insect remains alive, and at the end of that period commences to
take on a yellowish hue, the larvae of the parasite showing clearly
through the skin of the abdomen (fig. 8).
1909] Toxoptera graminum and its Parasites 81
Some studies were made of the final larval stage by Miss
Weeks, of Manhattan, Kansas, under my supervision, the results
of which were given in brief by me at the f
first meeting of this Society in Boston, / ‘
and are as follows:* lz7rm™)
A wingless 2 of T. graminum was NE AN
taken from the field when beginning to tees
take on the yellowish color, which first — ia
denotes parasitism, and which precedes on a
the amber color of the more advanced ‘Capaeetans aNy ‘
stage. Evidently this was on or about ar BR we
the sixth day from the time of Oviposi- [<= \ /
tion by the Lystphlebus, the larva being ) \
at the time quite full grown and occupy- _, P16. 8 Wingless female
5 : oe : 3 of spring grain-aphis, con-
ing its normal position in the posterior taining Jarva of the par-
bage en theabdemen of the host insect. ase | Fyswienus inc.
Much enlarged (original).
From all that we have been able to learn
the very young parasite larva takes this position, and, refraining
from moving about to any considerable degree, it does not come
in contact with or injure any vital organ of its host, the effect
being to reduce the period of reproduction of the 2 Toxoptera
to a very few days at most. But on reaching its full larval
development, it seems suddenly to become more active, and
with its first extended travels within the abdomen of its victim
the last spark of life remaining in the latter is extinguished. It
was just at this point that the observations herein recorded were
made.
On bringing the 2 Toxoptera from the field and placing it under
the microscope, the Lystphlebus larva was observed to work its
way actively about as shown by the accompanying plate, these
movements being always forward and easily observed through
the skin of the victim until this had become too opaque. The
movements of the parasite larva were accompanied, on the part of
the victim, by a clutching of the leaf and a rigidity of the muscles
of the limbs. But it has since been learned that this rigor mortts
has little if anything to do with the adherence of the cocoon to
the leaf surface, as was previously supposed by me, except per-
haps to temporarily hold the body of the aphid fast until the
parasite larva can anchor it more securely. (See Proc. Ent. Soc.
* These observations were afterwards published in Proc. Ent. Soc. Wash.,
Vol. IX, Nos. 1-4, pp. 110-118, Plate VI.
“OSVM WA
‘NHSV IOILIUG SOAATHAISAT NI AAOTHANA IVdNd AHL AO DNINOIHSVS AHL
tl ef @l
Gt % S
aa @
)
@):
Ly
a:
ney
DH
"AIX GLVTd ‘II “IOA ‘y ‘S ‘WT SIVNNY
1909] Toxoptera graminum and its Parasites 83
Wash., Vol. IX, p. 111.) Later observations by Mr. Kelly on the
movement of the parasitic larva, made from beneath and after
the dorsal skin of the host had become too opaque to continue
them from above, shows that the ventral skin of the Toxoptera
splits and the sht thus formed widens, as the skin assumes a ro-
tund shape, until it becomes nearly circular. The parasite larva
spins a cocoon of silken threads within the empty skin of its host,
and, by reason of this disc-like hole in the ventral surface of the
host skin, attaches the whole structure firmly to the surface of the
leaf.
Taking up these observations in detail, figure 1 of plate XIV
shows the host as when first placed under the microscope at 11
A. M., while she was still alive and at a time when she seemed to
tighten her grasp on the leaf with each movement of the parasite
larva within her body. Between 11 a. M, and 11:35 A. M. the
Lystphlebus larva had made three complete revolutions in the
body of its host, some of the different positions assumed, as also
the gradual shaping of the skin of the now dead Toxoptera being
illustrated by figs. 2-7 inthe plate. Between11:35 and 11:40 A.M.,
the larva had completed another revolution, probably the fourth
from the beginning. The fifth revolution was completed by
11:30 A. M., and the sixth at 11:58 a. M., while the seventh was
completed by 12:07 p. M. With the eighth revolution, shown by
figs. 8 and 9, completed at 12:11 p. M., the skin of the host insect
had been brought into its rotund shape and the larva had just
begun to contract preparatory to pupation. At 12:15 the ninth
EXPLANATION OF PLATE XIV.
Fic. 1—Position of larva of Lystphlebus tritici in body of wingless adult 92
of Toxoptera graminum, just before beginning its revolutions for fashioning the
body of the Toxoptera into a pupal envelope, 11 A. M.
Fics. 2—7.—Some of the positions assumed by the Lysiphlebus larva
between 11 a. M. and 11:35 a. m., during which time it made three complete
revolutions.
Fics. 8, 9.—Positions during and at completion of eighth revolution, 12:11
P.M.
Fic. 10.—Position at completion of ninth revolution, showing contraction
of the larva, 12.15 P. mM.
Fic. 11.—Position at 12:20 p. m.
Fic. 12.—Position at 12:22
Bie, 13.—Position at 12:27
Fic. 14.—Position at 12:32
Fic, 15.—Position at 12:32
84 Annals Entomological Society of America [WolsJEr
rotation (fig. ro) had been completed and the larva had decidedly
contracted. Fig. 11 shows the situation of both Jarva and pupal
envelope at 12:20; fig. Te) at 12:20) fic. 13,9ab 12:27 fo. 14, ab
12:32, and fig. 15, at 12:324 p.m. At 12:35 P. M. there were ap-
parent on the body of the parasite larva small, roundish cells of
a yellow color. By this time the skin of the host had reached
the semiglobular, typical form, which did not materially change
afterward, though a slight movement in the larva could be
detected up to 4 P. M., after which the covering had become too
opaque to permit of further observations. Thus, within the
space of five hours the living body of the host had been trans-
formed into a tough, dried, hardened protective covering the
cocoon of the parasite during its short pupal stage, by a process
of manipulation by the larva and a natural tanning process which
renders the skin of the dead Toxoptera so impervious to the
weather that these skins so prepared, may be washed off the leaves
by beating storms, yet do not become easily disintegrated, and
may often be found in quantities on the ground underneath the
infested grain plants.*
While the normal position of the larva of
the parasite in the abdomen of the host is indi-
cated in Plate 1, fig. 1, there are exceptions,
as shown by a case where, in a very young
Toxoptera, the position illustrated in text fig. 9
was assumed. This, however, is unusual. Again,
while the transformation of Lysiphlebus from
egg to adult takes place within the pupal
envelope, as previously described, and the adult
makes its way out by way of a small disc which Fic. 9 — Unus-
it gnaws through both cocoon and skin and beet SEG
pushes outward, as shown in fig. 10. Yet, ¢riticit in body of
sometimes, the larva works out of this case pre- ee ee eee
: ge
maturely, as shown in text fig. 11. This may, (original).
perhaps, be due to some mishaps in constructing
the anchorage previously mentioned. Mr. C. N. Ainslie found
several of these prematurely issued larve on the ground in
wheat fields in Kansas, under very seriously infested grain plants
where the Toxoptera were very much parasitized.
* Nore: Mr. Kelly has since found that the parasite slits the ventral skin
of the host, and, widening the rent spins a cocoon within the aphis skin and
anchors it fast where the slits are situated.
~
1909] Toxoptera graminum and its Parasites 85
The celerity with which an invasion of Toxoptera is overcome
by Lystphlebus, is frequently a matter of wonder, as it hardly
seems possible that this host alone could be the source of the
=a pal al (al Oa =—
appearance after a few warm days have
elapsed, in the midst of an unseason-
ably cool spring, preceded by a winter
abnormally mild.
In order to determine the origin of
these myriads of parasites, Messrs.
Kelly and Urbahns began a long series
of experiments at Wellington, Kansas,
to determine whether there might not
be a multiplicity of host species from
which great, numbers of Lystphlebus |
would emanate to fall upon and destroy 1
Toxoptera whenever it becomes exces-
sively abundant. The entire failure to L}_ ay Hel
introduce these parasites in advance of Fic. 10—Dead “green
; f ; bugs,’ showing hole from
an invasion of this character from the which the matured parasite
south, as was the case in Kansas in of Lystphlebus tritict emer-
May, 1907, has indicated that such intro- §o ya Stil ateched tes
ductions were not possible and that to pushed back; the bottom
: : 5 figure shown the parasite
attempt it was a veritable carrying Of emerging. Enlarged .. (ori-
coal to Newcastle. ginal.)
Female Lysiphlebus from Aphis brassice taken from the field
were first bred into Toxoptera from which adults were obtained.
swarms of individuals that make their THT
|
WM Fic. 11—a, Larva of Lystphlebus tritict working its way premature- [
ly from body of Toxoptera graminum; b, same larva, somewhat further
developed; c, same, still more developed. Enlarged (original).
86 Annals Entomological Society of America [Vol. II,
This experiment was several times repeated. Female Lysiphle-
bus reared under cover from A phis maidtradicts were in two cases
bred into Toxoptera and adults obtained. Female Lysztphlebus
were reared from Aplus setarte from the field and the issuing
parasites in two cases were bred into Toxoptera and adults
secured. This was also repeated several times. Female Lysi-
phlebus reared from Aphis maidis from the field, the adult par-
asites were bred into Toxoptera and adults emerged. Repeated
several times. Female Lysiphlebus from A phis gossypit from New
Mexico and the adults successfully bred into Toxoptera. Female
Lysiphlebus reared from Aphis setarie from the field were bred
into Toxoptera thence into A. maidiradicis and back into Toxop-
tera. Female Lystphlebus from A. setarie taken from the field
and bred into Yoxoptera, then back into A. setarie again into
Toxoptera and then into A. brassice from which adults were
secured.
In many cases these breedings were reversed. The only cases
of failures were in attempting to breed Lysiphlebus from Toxop-
tera into Chattophorus; and in attempting to breed Lysiphlebus
from 7 oxoptera into Macrostphum rudbeckie. These experiments
were reversed with the same results.
These experiments will be carried further another year, as it
seems that host habits if we may term them such may possibly
divide the specific position of the parasite in future.
The following diagram will serve to illustrate the additional
host insects of the species of Lystphlebus and A phelinus that are
known to also attack Toxoptera. This shows the varied host
sources from which parasites may come in case of a serious inva-
sion of Toxoptera.
2)
~“
1909| Toxoptera graminum and its Parasites
Toxortera
Schizoneura lanigera, S.C.
Myzus’ mahaleb, Ohio.
Pemphigus fraxinifolii, Kan.
Aphis monarde, Ohio.
Aphis sacchari, Java.
Siphonophora rosa, Fla.
Glyphina eragrostidis, Ind.
Tetraneura colophoidea, Md.
Aphsisi spie((2)), SiC
Aphis maidis, Iowa.
Myzus persece, Col.
Chaitophorus viminalis,Col.
Aphis gossypii(?), Col.
getarie, Kan.
-Aphis brassice, Kan.,Cal.
--Aphis maidiradicis, Kan.
---Aphis maidis, Kan.
gossypii, Kan,
BIBLIOGRAPHY.
(1) Rondini, Camillo, Nuovi Annali delle Scienze Natural, Serie III, Tom, VI,
pp. 10-12, 1852. See also Afidicidi, 1847.
(2) Passerini, Giovanni, gli Afidi. Pamphlet, p. 25, 1860.
(3) Horvath, G. Rovartani Lapok, Vol. I, p. 1438, 1884.
(4) Sajo, Karll, Zeitschr. f. Pflaugenkrankhaiten, Vol. IV, p. 4, 1894.
(5) Schouteden, H. Mem. Soc. Ent. Belgique, XII; p. 231, 1906.
(6) Sanderson, E. D., Bull. 46, Bu. Ent., U.S. Dept. Agr., p. 93, 1904.
OBSERVATIONS ON A GALL APHID (APHIS ATRIPLICIS L.).*
PauL HAYHURST.
The subject of this article is the common greenish insect of
the genus A plis which colonizes the dorsal surface of the leaves
of the white goosefoot or lamb’s quarters, Chenopodium album,
and the nearly allied orache, Atriplex patula, in summer. The
margins of the infested portion of the leaf always curl up longi-
tudinally and meet above the midrib, forming an imperfectly
closed tube.
According to the determinations of Mr. T. A. Williams and of
Doctor Forbes, A phts atriplicis also attacks the leaves of culti-
vated beets in Nebraska and Illinois. This would not be at all
surprising, since the genus Geta is closely related to Chenopodium
and Atriplex. Doctor Bruner wrote me that Mr. Williams was
well acquainted with the insect and knew its characteristic habits
on goosefoot. This aphid is very abundant on the common
goosefoot and orache, from which it might easily become trouble-
some on beets.
At Fredonia, N. Y., it was everywhere abundant on the white
goosefoot during August, 1908, when I first became familiar with
its habits, although I did not determine it specifically until the
following January. Most plants of this host were badly infested
both in the vicinity of Fredonia and also at Lily Dale, Jamestown,
and Chautauqua, N. Y. It has been reported as infesting this
plant from Missouri, Kansas, Minnesota, Colorado, Hlinois and
Nebraska. Professor G. Del Guercio, Florence, Italy, writes me
that this species is everywhere common in Europe on plants of
the goosefoot family. I find in the literature that it has been
reported from the following countries: Sweden, Germany, Italy,
England and Belgium.
In habits this species is unique. The insects were infesting
only the dorsal surface of the leaves and no other part of the
plant when I was studying them last August. They were clus-
tered chiefly along the midribs and main veins. The elongated
galls (Fig. 1), which were formed by the sucking on the upper sur-
face, partially protected the occupants from rain and sun. The
margins of the leaves were never rolled in or convoluted more than
I have shown in Fig. 1, a—b, which are cross sections of represen-
*Contributions from the Entomological Department of Bussey Institution,
Harvard University, No. 5.
88
1909] Observations on a Gall Aphid 89
tative galls. Thus whenever the galls retained the normal posi-
tion of the leaves with the dorsal surface upwards, they were easily
filled with rain-water; in most cases, however, they were partly
or entirely reversed bringing the ventral surface upwards and
directing the long aperture of the gall downwards. Such galls
were effectual water-sheds and the inhabitants were always dry,
while those that were not reversed carried more or less water
mixed with the excretions and exuvia of the plant lice. This
liquid was evidently detrimental to the insects, many of which
were killed by a parasitic fungus. An ample supply of these
brownish dead aphids was sent to Professor Roland Thaxter of
Harvard University, who replied as follows: ‘‘* * * * the fungus
which has attacked them is Entomophthora aphidis, a universally
distributed form with which two others are often associated. In
your material, however, I see but the single species mentioned.”’
It is clear, then, that the reversal of the gall is a decided advan-
tage to the species. I was not able to determine how this reversal
was brought about. It was evidently one of the curious effects
on the leaf caused by the sucking of the insects, and may have
been developed by natural selection for their protection from
dampness and disease. The upturned whitish under surface of the
leaf-galls rendered the infested plants very conspicuous, and gave
them the deceptive appearance of being attacked by a downy
mildew. I have, in fact, noticed this phenomenon in the Cheno-
podiums in Chautauqua County, N. Y., for several years and had
always supposed it to be the work of a fungus until I examined
them critically last summer.
A black aphid, the Aphis rumicis L. of authors, commonly
infests the lower surface of the leaves and the inflorescence of these
plants. It also sometimes occurs mixed in the colonies of the
green aphids under discussion. This habit undoubtedly accounts
for Buckton’s mixed descriptions. The green and the black
forms are entirely distinct species easily separable by pronounced
structural differences, as pointed out in the descriptions farther on.
During the first week in August, 1908, the winged vivipare
(viviparous females) of Aphis atriplicis were leaving the galls in
great numbers, a general migration being apparently at its
height. After the middle of August pupee were not common in
the galls, which were almost deserted, except for a few wingless
vivipare and larvee. Connold, 1902, observed a similar migration
in August in England. I failed to find what became of the winged
go Annals Entomological Society of America [Vol. IT,
females. Many individuals were found isolated on bean leaves,
but none of these succeeded in establishing colonies. The mi-
grants were certainly not depositing larvae on the normal leaves
of the Chenopodiums. About the middle of October, 1908, at
Forest Hills, Mass. I found many small plants of this genus grow-
ing in soil that had been cultivated or newly made after July.
These plants must have come up lateinsummer. They were cov-
ered with the galls of thisspecies. It is evident, then, that they were
colonized in August. Undoubtedly the winged females observed
migrating at Fredonia in the same latitude were merely colonizers
on tender young Chenopodiums starting up in damp situations.
It is highly improbable that these were true migrants going to
some distinct host. As a rule the sexes of truly migrating plant
lice develop on some woody plant called the primary host, while
the summer forms of the species live on a succulent herb, the
secondary host. Since the sexes of Aphis atriplicis developed in
great numbers on the Chenopodiums last fall, they are undoubt-
edly confined in their life-cycle to these plants and their allies.
On September 21st the perfect sexes were first noticed on the
white goosefoot at Forest Hills. Small, lank, wingless males of
a yellowish color, with blackish appendages, were actively climb-
ing about over the large clusters of plump oviparous females,
and copulation was observed as late as the 30th of October.
Large numbers of these females were found abundantly on the
goosefoot plants in this neighborhood. The males were few in
number, between 5 and ro per cent. of the total number of indi-
viduals (by guess). But they were making up for their fewness
by great activity, and copulation was often seen. Most of the
females had left the summer galls and were on the seed-heads,
where they were frequently mixed with black aphids, the so-
called Aphis rumicis. Only a small proportion of the individuals
were found in the galls during the periods of oviposition. No
viviparee were seen at Forest Hills, and of many specimens taken
throughout the fall, none proved to be viviparous on examination
in the laboratory. Viviparity doubtless ceases or becomes
exceptional in this species with the development of the perfect
sexes. There is evidently a general migration of these sexes or
of the sexupare producing them from the galls to the seed-heads
in the fall, where the eggs are usually laid. Oviposition was
observed during the entire month of October. The eggs were at
first light yellow in color, then deep green, finally becoming shin-
1909] Observations on a Gall Aphid QI
ing black a few days after they were laid. They were generally
placed irregularly on the calyces and seed capsules and on the
small leaves of the upper branches. I examined many of the
large galls finding some eggs in them, but not nearly so many as
on the seed-heads. On October 21st I did find many eggs in
some frosted wilted leaves. The general instinct of the females
was plainly to carry the egg-laying as far up towards the seed-
bearing portion as possible. On Nov. 11th no more living aphids
of this species were found, for a freeze had killed all the Cheno-
podiums.
This habit of ovipositing on or close to the seeds is an instruc-
tive adaptation to the nature of the host. | The winds and storms
of the winter of rg09 have broken off all the seed-heads and leaves
of the Chenopodiums, and at the time of writing (April) even the
bare stalks of the plants rarely remain standing. The seeds have
been distributed far and wide and many of the eggs of the aphids
must have been carried with them. The young stem-mothers
on hatching will be able to find their natural food easily avail-
able. Were most of the eggs laid on the leaves, they would be
so scattered by the winds that very few of the larve could find
suitable food in the spring. Since most species of Chenopodium
and Atriplex are annual, the stem-mothers would suffer the same
mishap if the eggs from which they hatched had been laid on the
ground beneath the host. In October and November, 1905, at
Columbia, Missouri I observed the large red Macrosiphum rud-
beckie Fitch ovipositing in great numbers on the debris under one
of its hosts, a perennial goldenrod. In this case new shoots would
have come up in the spring around the old plants, and the larvee
on hatching would have found ready food. Here then, there was
no necessity of the mothers entrusting their progeny to the
capriciousness of the windy winter elements.
The oviposition of our goosefoot aphid in the seed-heads
explains its world-wide distribution. Professor M. L. Fernald,
Harvard University, has written to me that Chenopodium album
is known to be an introduced plant from Europe, whence it has
followed the trail of the early American settlers. In Professor
Fernald’s own explorations in the forests of Maine, New Bruns-
wick and Quebec this plant was never found except about settle-
ments. Atriplex patula is likewise recognized as an introduced
species. These plants are both common European garden weeds
whose seeds, with the eggs of their insect foes, could easily be in-
troduced with any rubbish into a new country.
92 Annals Entomological Society of America (Vol, Ef,
The wingless males are interesting types of degeneration
(Fig. 3). They have not only lost their wings, but the thoracic
lobes are also disappearing. In most specimens of about a score
examined these structures are distinctly traceable, although re-
duced, and compare well with those of any winged form of the
genus Aphis; but in some individuals they are degraded to mere
vestiges in the shape of irregular dusky marks. In these more
degenerate specimens only the ocelli, present in all winged aphids,
prove that they were, until comparatively recently, winged
insects. In this species we may hope to find occasional males
still retaining their wings. Weed has shown that both winged
and wingless males occur in Cladobius salicis L., and Gillette
has found both forms in Aphis torticauda Gillette. No winged
males of Aphis atriplicis were found this fall, although I kept this
possibility constantly in view. On October 15th I did find three
winged males in copulation with females of this species, but the
males could not possibly be the same species, for they were gener-
ically distinct. This observation suggests the possibility of hy-
bridization in the plant lice. Weed in 1891 observed the copu-
lation of a male of Cladobius salictt Harris with a female of Lachnus
platanicola Riley. These records also teach us that the mere pairing
of two plant lice does not prove their specific or generic identity.
The former can, however, be safely inferred when many males
are found pairing with a large number of females possessing
similar structural characters.
The wingless males of a red Myzus (?) on cherry (not Myzus
cerast Fab.) collected at Forest Hills last fall, have degenerated
much farther than those of our Chenopodium aphtd. Here all
traces of thoracic lobes have disappeared, and the ocelli are scarce-
ly discernible. A still farther step in degeneration has apparently
taken place in the wingless male of the corn-root aphis (A phis
matdi-radicis Forbes), if we may judge from Forbes’ figure and
the descriptions by the same writer, Weed, and Davis, in which
no mention is made of the ocelli. There would be no way of recog-
nizing this male as a descendant of a winged male ancestor were it
not for these intermediate forms, which I have discovered. This
process of reduction of structures correlated with the organs of
flight is an instructive illustration of what must have occurred long
before in the evolution of the wingless vivipara, which has even
lost all traces of ocelli. These facts also suggest the probability
that this female in the Aphidide is not a case of paedogenesis as
1909] Observations on a Gall Aphid 93
usually supposed, but is morphologically equivalent to a winged
vivipara which has lost her organs of flight and all correlated
structures through disuse. Living as parasitic insects in the
midst of a luxuriant supply of food all superfluous structures have
been dispensed with according to the well known laws of para-
sitism. When the males of a species are wingless, they must pair
with females which have developed on the same plant and which
may be of the same descent. The general migration of A phis
atriplicis in August observed by Connold and myself undoubtedly
tends to counteract cross fertilization, since the bisexual colonies
on any plant in the fall would be of mixed descent. According
to the literature which I have seen, wingless males are known in the
following species: Pemphigus attenuatus Osborn, Schizoneura cornt
Fab., S. americana Riley, S. lanigera Hausm., Hamamelistes
spinosus Shim., Hormaphis hamamelidis Osten Sacken, Chermes
and Phylloxera, all species as far as known, Melanoxanthus salicis
L., Lachnus nudus De G., L. piceicola Cholodk., Aphis maidt-
radicis Forbes, A. atriplicis L., A. malt Fab., A. carbocolor Gil-
lette, A. torticauda Gillette. Buckton reports wingless males in
ten other species, but since I know the winged males in four of
these and since his work seems so generally superficial, I do not
give his list.
Aphis atriplicis was greatly parasitized both at Fredonia and
at Forest Hills. On some plants nearly all the aphids were
killed. I am indebted to Mr. C. T. Brues, Curator, Public Mu-
seum of the City of Milwaukee, Milwaukee, Wis., for the deter-
mination of the following parasites which I reared from this spe-
cies: Lystphlebus eragrostaphidis Ashm., Fredonia, N. Y., Aug. 3,
1908. Several specimens. From viviparous aphids. The other
parasites were all reared from Forest Hills material and must have
developed very largely in the oviparous insects. Lysiphlebus
eragrostaphidis Ashm., Oct., to, 23 and 30; many specimens.
Pachyneuron micans Howard. Oct. 30, one specimen. Asaphes
rufipes Brues, n. sp. Oct. 30, four specimens. Aphycus ? sp. an
Encyrtid. Oct. 30, one specimen. Figites, sens. lat., aberrant
Cympide. Mr. William Beutenmuller suggests that these may
belong to the genus, Allobia, known to be parasitic in plant lice,
but no one in this country has time to study them for me. Oct.
30, nine specimens.
Several lady beetles were collected from galls of this aphid at
Fredonia. Adult examples of these were kindly determined for
94 Annals Entomological Soctety of America [Vol it,
me by Mr. E. A. Schwarz through the courtesy of Doctor Howard
as Scymnus americanus Muls. and Hippodamia 13-punctata L.
The aphids were not attended by ants. This may have been
due to the dry time of the year, when, as Professor Wheeler tells
me, these insects are not generally so active outside of their gal-
leries. It is possible, however, that Aphis atriplicis does not
attract ants at any time of the year. Mr. J. J. Davis has found
that Szpha flava Forbes is never attended by ants in Illinois and
I have noticed the same fact relating to this species in the early
summer in Virginia and New York when ants were plentiful in
the colonies of other aphids. It is also a matter of common
observation that woolly aphids are rarely or never cared for by
ants. ;
The identification of this species was not difficult on account
of its peculiar habits. The original description by Linnzeus is as
follows: ‘‘Habitat in Atriplicis littoralis foliis, quae inde revolun-
tur longitudinaliter in cylindrum, intra quem vagantur obtecte.
Deser. Corpus viride: corniculs minutis. Oculi nigri.” It will
be seen that this description is less ambiguous than those of
many species unhesitatingly referred to Linnzus, and I prefer
to use 1t to avoid multiplying names. Specimens of the winged
and wingless viviparee were sent to Professor Del Guercio who is
familiar with the species in Europe, and he found them to be
identical. He placed this species in his genus Uraplis which is
distinguished from other subdivisions of Aphis by the cornicles
being shorter than the cauda. Since the cornicles are distinctly
incrassate, he proposed a new subgenus of Uraphts, to be called
Hayhurstia Del G. This subgenus would then include all species
of A phis with incrassate cornicles which are shorter than the cauda.
Del Guercio separates Siphocoryne Pass. from Aphis by the cor-
nicles which are ‘‘more or less long, clavate.’’ (Nuov. Relazioni
R. Stazione di Ent. Agrar. Firenze (1) No. 2, 1900, p. 142). By
general consent those species of Aphis with more or less clavate
cornicles which are longer than the cauda have been put in the
poorly defined genus, Szphocoryne. In other words, Hayhurstia
Del G. is separated from Szphocoryne Pass. merely by the length
of the cornicles and a supposed difference between clavate and 1n-
crassate. ‘These I consider too indefinite to be reliable generic
characters. When forms are found with the cornicles about the
length of the cauda, very careful measurements must be taken to
determine the genus, and ocular micrometers are not always at
1909} Observations on a Gall Aphid 95
hand. Since every gradation exists in the degree of the bulge in
the cornicles of the genus Szphocoryne from forms like S. saliczs
Monell down to S. avenae Fab. which is scarcely incrassate at
all and that on the inner side, I cannot recognize either Szpho-
coryne or Hayhurstia as valid genera, but would rank them both
as subgenera of the genus Aphis. I therefore consider the follow-
ing grouping of those species of Aphis and Siphocoryne with a dis-
tinct cauda worth testing. This is modified from Del Guercio’s
revision of Aphis (Redia, Firenze Vol. 4, Fasc. 1, 1906, pp. 191-2.)
1. Cornicles longer than cauda.
a. Cornicles cylindrical or gradually attenuated from base.
Subgenus Aphis L. (Fig. 13.)
Aphis sambuci L.
b. Cornicles clavate or incrassate at or beyond base.
Subgenus Siphocoryne Pass. (Fig. 11.)
Aphis nymphee L.
2. Cornicles equal to cauda or shorter.
a. Cornicles cylindrical or gradually attenuated from base.
1. Cornicles shorter than style.
Subgenus Uraphis Del G.
Aphis genistz Koch.
2. Cornicles equal to style.
Subgenus Microsiphon Del G.
Aphis tarmentillz Pass.
b. Cornicles incrassate.
N. Subgenus Hayhurstia Del G. (Fig. 12.)
Aphis atriplicis L.
Aphis atriplicis L. seems never to have been carefully
described. In the following descriptions all colors were taken
from the living insects with a high power hand lens, daylight,
and many individuals were compared. All measurements of the
bodies are from formalin material and are therefore reliable, while
those of the appendages are from balsam mounts; the lengths
given are averages of six or more representative specimens. Fig-
ures in parentheses are extremes. Del Guercio states that the
colors especially of the wingless vivipare in Italy vary greatly
according to the host and other conditions so that ‘“‘now greenish-
yellow forms preponderate, and now those of a yellowish-green,
green, olive green and olive color, sometimes, indeed, so deep in
tint as to seem brownish with greenish reflections.”
The lobes of the mesothorax are figured for clearness, because
they are not generally mentioned individually by authors (Fig. 2).
Their color is here treated as of specific value, but their form is
not described for this purpose. Their structural details present
marked constant differences, which have apparently never been
carefully studied. They are already called the prescutum, scu-
96 Annals Entomological Society of America [Vol 11,
tum, (scutal lobes), scutellum and postscutellum by some aphid-
ologists, and I accordingly use the same terms, although I do not
know their homologies. I avoid the long term viviparous female
giving preference to its simple equivalent, vivipara (ple),
which, being a classical word of the feminine gender, means liter-
ally the viviparous female. Since all viviparous plant lice are
necessarily females, I can see no ambiguity in the use of this word.
Winged vivipara. Head dusky green; antenne dusky with pale
articulations, sparsely hairy, joint III, usual number of sensoria
9 or 10, sometimes 11-13 (Figs. 10, 15, 18); eyes dark red; beak extend-
ing beyond transverse sternal suture, but not reaching middle coxe.
Pronotum dusky green; lateral tubercles present, not prominent, longer
than broad. Mesothorax greenish yellow; prescutum, scutal lobes
and postscutellum blackish, scutellum (in formalin) brownish with
black anterior and posterior margins; mesosternal plates brown, meta-
sternal region dusky; Ist and 2nd discoidal veins of the forewing more
or less distinctly joined to radius, never completely atrophied at base
(Fig. 6). Base of femora brownish yellow, elsewhere dusky; tibiz
brownish yellow with dusky apex, tarsi dusky to black. Abdomen
green, varying to yellowish green. Dorsum with irregular, variable,
dusky maculations, most commonly resembling Fig. 5. The four
large lateral spots in front of the cornicles distinct, dusky to blackish.
The seven small, lateral, obtuse tubercles distinct but not conspicuous,
mostly broader than long (Fig. 5). Cornicles dusky, swollen at and
beyond the middle, a little shorter than hind tarsus, surface smooth,
never imbricated (Figs. 8, 12); cauda uniformly pale yellowish from
base to apex, the margins black anterior to the distinct median constric-
tion, posterior to which are 3 pairs of long curved sete; anal plate beset
with several similar sete, 6-8 along posterior margin.
Measurements. (Antennz measured from base of III to apex of
filament). Length of body, head to base of tail, 149 mm. (1.39-
1.70); greatest width of abdomen, .662 (.61-.696). Antenne, .97—
1.00 TIL, 33531V, 1552 Ve lobe Vi0 (Geape) 107, filament,.25- Bore=
wing, 2.0-2.26.. Cornicles, 0.12; cauda, 0.155.
Pupa. Color similar to the winged vivipara, with indistinct or no
maculations except irregular deeper green marks on mesothorax;
wing-pads dusky.
Wingless vivipara. Head dusky yellow with two median dusky
spots close together. Antenne I-II concolorous, elsewhere light
brownish, the distal joints becoming dusky; without sensoria except
the usual one at apex of V and the group at the apex of the scape of VI.
Eyes reddish black or black. Beak extending to or a little beyond
mesocoxe. Thorax yellowish green varying to green; a dusky dorso-
lateral impressed spot on each segment, the one on the mesothorax
longitudinal linear, often broken into a row of small spots; thorax
otherwise without maculations. Distinct prothoracic lateral tubercle,
longer than broad. Legs yellowish-brown except the dusky apex of
tibiz and the tarsi. Abdomen concolorous with thorax, without macu-
1909] Observations on a Gall Aphid 97
lations except 4 or 5 small dusky spots forming a dorso-lateral longi-
tudinal row continuous with the thoracic spots, the last distinct spot
on about the 5th segment; one indistinct spot just mesal to the cornicle,
often absent. Cornicles yellowish with apex dusky, the entire appen-
dage usually not dusky. Whole body lightly pulverulent. Other-
wise as in winged vivipara.
Measurements—Length of body, 1.54 mm. (1.35-1.70), width,
:009 > (.022—.690). — Antenne: 084 (.516—.705); ITI .20, IV, .077, V
09, VI (scape) .077, filament .142. Cornicles, .114, cauda, .172.
Ovitpara. This form can be easily told with a hand-lens from the
wingless vivipara by the dusky incrassate hind tibie with many
sensoria and the blackish vaginal plate. Detailed description as fol-
lows: Eyes black, rarely reddish black. Legs dusky except the yel-
lowish—brown base of femora and greater proximal portion of fore and
middle tibie, hind tibie dusky, proximal half swollen, 30-40 nearly
circular sensoria on both anterior and posterior aspects (Fig. 4). Cor-
nicles dusky: Cauda pale yellowish distally, or entirely dusky; often
not constricted. Otherwise similar to the wingless vivipara.
Measurements. Length of body, 1.45 mm. (1.24-1.70), width
065 (.478-.652). Antenne .553 (.516—.671); III .129, IV .06, V
.0774, VI (scape) .077, filament, .122. Cornicles, .112, cauda, .146.
Wingless male. Head and antenne dusky to blackish. Antenne
(Fig. 17) with average number of sensoria as follows (extremes in
parentheses): joint III, 14-15 (11-24); IV, 9-10 (7-14); V, 10-11
(10-14); VI, usual group at apex of scape, often with a single sensor-
ium near its middle. Eyes blackish red; ocelli conspicuous.
Beak extending just beyond mesocoxe. Pronotum dusky, lateral
tubercle distinct, mesothoracic lobes brownish to dusky, traceable as
in Fig. 3; in several specimens they are reduced to irregular dusky
marks. Legs dusky to blackish. Abdomen dusky yellow, very varia-
able in the tint of yellow, dorsum with irregular, variable dusky marks,
usually resembling Fig. 3. Lateral tubercles relatively more pro-
nounced than in the females. Cornicles dusky. Cauda dusky, not
constricted (Fig. 19). Otherwise as in winged vivipara.
Measurements. Length of body, 1 mm. (.957—1.04), width, .481
(4380-500). Antenne .39) (72=1.02) IE, 258: IV, 142° V, .149,
VI (scape) .094, filament, .215. Cornicles, .066; cauda, .113.
Eggs, oval, shining black, .60 x .327 mm. Dissection of many
ovipare collected Oct. 9th, showed that each individual contained
from one to eight eggs. The number evidently depended on the size
as well as the age, since some large specimens were opened that
contained no eggs. The usual number was two to six.
-The common black aphid infesting the Chenopodiums, as stated
above, is entirely distinct from Aphis atriplicis L. For the sake of
comparison I have drawn the principal systematic differences which
may be summarized as follows:
Winged vivipara—Antenne with many (fully 15-16) sensoria on
joint III; 4-5 on IV (Fig. 16). Basal half of costal vein of fore-wing
strongly bent anteriorly, bases of Ist and 2nd discoidals always com-
98 Annals Entomological Society of America [Vol. IT,
pletely atrophied. Cornicles (Figs. 9, 13), never swollen, but broad
at base tapering gradually to apex, distinctly imbricated, black, about
1.5 times length of hind tarsus. Cauda (Fig. 14), stouter, apex more
rounded, beset with many stout curved setz, bicolored, the basal half
between the black margins being pale yellowish, while the distal half
is dusky to blackish. The prothoracic and 2nd and last abdominal
tubercles very pronounced, longer than broad.
Measurements. Length of body, 2 mm. width .98; antenne
1.25; wing 2.75; cornicles, .20; cauda, 1.5 (averages from several
specimens. )
The life history of this black aphid is different from the
green species. It never forms galls, usually infests the under
side of the leaves or the inflorescence in summer as well as fall,
attacks Rumex and other weeds besides the Chenopodiacee; and
migrates in the fall from these plants to the wahoo (Luonymus
atropurpurea.) I observed this migration to the wahoo last fall
at Forest Hills where the so-called sexuparee were depositing ovi-
parous larve on the under side of the leaves. These true mi-
grants were precisely identical in structure with the winged vivi-
paree on Chenopodium.
Osborn established the migration of this wahoo aphid (A phis
rumicis ?L. = A. euonymi Fab.) in Iowa in 1894. Either this
species or a closely similar one migrates between the same hosts
im Europe.
It is my privilege to acknowledge Professor Wheeler’s kind
criticisms in the preparation of this article.
LITERATURE
Aphis atriplicis Fab., Fabricius, Syst. Ent., 1775, p. 737, No. 21, is a distinct
species belonging to the Aphis rumicis group.
Aphis atriplicis L., Linnzus, Fauna Svecica (ed. altera), 1761, p. 262, No. 1000.
Original description. Sweden. Atriplex littoralis.
—chenopodii Schr., Schrank, Fauna Boica, Vol. 2, Pt. 1, 1801, p. 109, No.
1196. Bavaria, Atriplex patula; descriptions of winged and wingless
vivipare.
Kaltenbach, J. H., Monogr. d. Pflanzenl., 1843, pp. 107-108. Germany. Cheno-
podium album, Atriplex patula. Descriptions of winged and wingless
vivipare.
Passerini, G., Gli Afidi, 1860, p. 31. Italy.
Ib. Flora’ degli Afidi Ital., in Bull. Ent. Soc. Ital. 3, 1871, pp. 151, 155.
On Atriplex patula and Chenopodium album.
Kaltenbach, J. H., Pflanzenfeinde, 1874, pp. 505, 508. In Germany. Same
hosts and habits. Sometimes affects all the leaves of the plants mak-
ing the leaves appear yellowed and sickly.
Buckton, G. B., British Aphides, Vol. 2, 1879, pp. 87-90, col. pl. 65, figs. 4-7.
All descriptions and figures apply to an insect of the Aphis rumicis type
except possibly the wingless male and the ovipara which apparently
belong to Aphis atriplis L. Notes on the galls and oviposition in the
galls. Atriplex sp. England.
Monell, J., Bull. U. S. Geol. & Geogr. Surv. Terr. Vol. 5, No. 1, 1879, p. 25.
Common on Chenopodiums at St. Louis, Mo., and at Carbondale, II.
(Thomas, C. 8th report Ins. Ill., 1879, p. 193.)
1909] Observations on a Gall Aphid 99
Lichtenstein, J., Les Pucerons, 1884, Chap. 4 (La Flore des Aphidiens) pp. 80,
85. Atriplex and Chenopodium.
Oestlund, O. W., Ann. Rept. Geol. & Nat. Hist. Surv. Minn., 14, 1886, p. 47.
Minnesota, Chenopodium album. (Ib., Bull. 4, Geol. & Nat. Hist.
Surv. Minn., 1887, p. 69.)
Bruner, L., U.S. Dept. Agric. Bur. Ent. Bull. 23 (0. s.) 1891, p.17. On the beet
at Ashland, Nebr. (T. A. Williams.)
Williams, T. A., Univ. Nebr., Dept. Ent:., Special Bull. 1., 1891, pp. 6, 10, 12,
20. Nebr. On cultivated beet, Rumex altissimus, Chenopodium album
and hydridum.
—chenopodii Cowen., Cowen, J. H., Colo. A. E.S. Bull. 31 (Tech. Ser.
No. 1), 1895, p.119. Descriptions of winged and wingless vivipare and
galls. Colo., Chenopodium album; galls collected in July.
Del Guercio, G., Nuov. Relazioni R. Staz. Ent. Agrar. Firenze (1) No. 2, 1900,
pp. 124, 132. Italy. Differential characters in table of species of
Aphis. Chenopodium album and Atriplex patula.
Forbes, S. A., Bull. 60, Ill., A. E. S.1900, p. 480. Illinois. Eggs in dried rolled
leaves of Atriplex. Suggests that this species may be identical with
Aphis gossypii Glover. (I do not think this possible. The two species
have totally different cornicles, those of Aphis gossypii are much longer
than the 4th tarsus, incrassate at the base, elsewhere cylindrical, imbri-
cated, black.)
Hunter, W. D., Bull. 60, lowa A. E.S., 1901, p.93. Partial bibliography. Hosts,
distribution.
Connold, E. T., British Vegetable Galls, 1902, pp. 233-4, pl. 101. England.
Atriplex augustifolia and patula. Abundant along the coast and
inland. Winged vivipare emerge in August. All stages including
sexes and ova in galls during the fall. (Notes on the insects show that
he evidently had colonies mixed with Aphis rumicis (?). Photograph
of infested plants showing galls.
Bignell, G. C., Ann. Rept., 72 Roy. Cornwall Polytech. Soc. 1904, p. 24 (Separate)
England. Atriplex latifolia.
Luff, W. A., Guernsey Soc. Nat. Sci., Rept. and Trans. for 1905, p. 88. Mention.
Schouteden, H., Cat. d. Aphides de Belgique. in Mem. Soc. Ent. Belg. 12, p. 216.
Belgium; Atriplex, Chenopodium album.
Sanborn, C. E., Kansas Univ., Sci. Bull. Vol. 3, No. 8, 1906, pp. 232, 235, 248,
254, 255. Compiled list of hosts.
PLATE XV.
Fic. 1. Leaf-gall; a, b, cross-sections.
Fic. 2. Winged vivipara. pt., pronotum; /t., lateral tubercle; p., prescutum;
s., scutal lobes; sl., scutellum; ps., post-scutellum ; mt., metatergite.
Fic. 3. Wingless male.
Fic. 4. Ovipara, hind tibia.
Fic. 5. Winged vivipara.
Fic. 6. Forewing of same.
Fic. 7. Forewing of Aphis rumicts (?).
Fic. 8. Cornicle of A phts atriplicis, distal end.
Fic. 9. Cornicle of Aphis rumicis (?), distal end.
Fic. 10. Third antennal joints of Aphis atriplicis, winged vivipara, from same
individual.
Fic. 11. Cornicles, two variations in the bulge in subgenus Szphocoryne.
Fic. 12. Cornicle of Aphis atriplicts.
Fic. 13.. Cornicle of Aphts rumicis (?).
Fic. 14. Cauda of Aphis rumicts (?).
Fic. 15. Third antennal joints of Aphis atriplicis, from four individuals.
Fic. 16. Antenna of Aphis rumicis (?), winged vivipara.
Fic. 17. Antenna of wingless male of Aphis atriplicis. (Thickness somewhat
exaggerated.)
Fic. 18. Antenna of winged vivipara of A phis atriplicts.
Fic. 19. Cauda of wingless male of A phts atriplicis.
All figures except 1 and 2 made with camera lucida.
ANNALS E. S. A. VoL. II, PLATE XV.
Pauli Hawhurst ded.
HOMOLOGIES OF THE WING VEINS OF THE APHIDID4,
PSYLLIDA, ALEURODIDAE, AND COCCIDA.*
EpitH M. Patcu.
INTRODUCTION AND GENERAL CONSIDERATIONS.
This study of wing venation was undertaken in the hope
that it might, for the one small group of Homopterous insects
it concerns, supplement in some shght measure other work in
wing-vein homologies which has been so vital a problem for insect
phylogeny.
The incentive to this study is to be found in ‘“‘The Wings of
Insects’? by Comstock and Needham. The choice of the parti-
cular group here considered was due to a personal interest in
aphids and to the fact that the homologies of the wing-veins
of this family were at that time practically untouched.
From the first the work has been under the direction of the
Department of Entomology, Cornell University and it is a pleasure
to acknowledge the kindness of Professor Comstock during my
association with this department as a student, and the patient
aid and constant interest and sympathy in my task of Doctor
MacGillivray.
To Professor Oestlund at the University of Minnesota is due
thanks for the determination of certain species of aphids which are
included in this study and for the personal interest with which
he has anticipated the results.
Although the work has been entirely under the supervision
of the Department of Entomology, Cornell University, the great
bulk of the data has been accumulated at Orono, Maine, instead
of Ithaca, New York. Several months of each winter during the
grogress of the problem, however, have been spent at Cornell in
revising data and in such consultation as has insured against
certain misinterpretations and other errors due to inexperience
in this type of work. At such times the essential points have been
in so far as is possible with the greenhouse aphids available,
verified while 7m residence in Ithaca, and the work with the Coc-
cide and Aleurodide has been done in the Entomological labor-
atory at Cornell University.
Since 1903 each season’s collection of aphids in Maine with
detailed notes has given me a very fair idea of where to secure
* A contribution from the Entomological Laboratory of Cornell University,
and Papers from the Maine Agricultural Experiment Station; Entomology No. 33
IOI
102 Annals Entomological Society of America [Vol ait.
given species and upon what plants at any time during the entire
season. This has given me a range of from g5 to 105 available
species belonging to at least 16 different genera and representing
all the types of venation in the entire family.
At the Maine Station two insectaries (one an unheated build-
ing of use only during the summer, and one a hot house), have
both been available for the segregation of colonies of aphids taken
previous to the development of wing pads in order that they could
be used at exactly the right time. This made possible a large
amount of material safe from the depredations of predaceous
insects and parasites, a condition which could never’ be relied
upon with chance collections in the open.
In view of the fact that for the work in hand, no specimen
could be used for the study of wing tracheation except during the
first few minutes after emergence from the last molt, and that
the nymphs could be studied only during a very limited time
before the developing wings became much folded in the sac,
and that some species gave conspicuously better results than
others by virtue of such reasons as the shghtly different angle
at which the wing pad of certain species are held, or to differences
in color, etc. :—it will be evident that the conditions outlined have
been most propitious for the study of this particular problem
with aphids.
The reasons for approaching the homologies of the wing veins
of insects by a study of the trachez that precede the veins were
so fully set forth and their validity so thoroughly demonstrated
by Comstock—Needham* ten years ago that this phase of the
question has long been too familiar to call for general discussion
here:
However, with each new group of insects studied in this way
conditions exist which may have a special bearing on the subject.
For instance, an ontogenetic study of the wings of certain insects
is not of any value in determining the homologies of the veins{
in which case entire dependence must be placed on a careful com-
parison of the veins of the mature wing.
On the other hand with wings so highly specialized by reduc-
tion of veins as those of the Homopterous group with which this
paper is concerned, the subject is a hopeless one to approach from
* The Wings of Insects. Amer. Nat. XXXII and XXXIII, 1898 and 1899.
t MacGillivray, A. D., Wings of the Tenthredinoidea 1906. Proceedings
of the Nat. Museum. Vol. XXIX, page 574.
1909} Homologies of Wing Veins 103
the comparison of the veins themselves. What chance is there
on the basis of the mature wings alone to homologize except by
the merest guess the veins of the Aleurodide with those of the
Psyllidee?
Yet fortunately and, in many instances, to my utmost sur-
prise, it was found that practically the whole story of the vena-
tion of the group of four families here concerned lies revealed
in the preceding tracheation and no one who follows that story
closely can fail to recognize in the two surviving veins of Aleurodes
for instance the radial sector and cubitus. (See page 122).
One of the simplest but prettiest and most clearly cut of the
demonstrations of the value of the tracheation in this connection
is to be found by comparing the venation and tracheation of
Chermes which shows at once that it is the wavering and unstable
R, which is lacking and not M as has heretofore been considered.
(See page 111). And the vagaries of R, in the whole group form a
fascinating study by themselves.
But exciting and interesting as is the story the trachez trace,
the quest for the homologies of the wing veins of aphids has lain
along a path tedious in many respects and beset with many diff-
culties.
The tracheze of aphid wings are very delicate, and when filled
with the medium in which the wing is mounted they become
invisible, so that many of the examinations and sketches made
had to be finished with rapidity, and often many mounts pre-
pared to verify a single point.
The wings studied in connection with this problem were pre-
pared in two ways. Part of them were mounted in glycerine
jelly after the method described by Comstock—Needham* and
part of them were mounted in xylene damar. With wings so
small and delicate as the aphids wings the latter was found in the
main more satisfactory. It took the balsam less time to penetrate
and render the trachee invisible which would have been an objec-
*“In making mounts of this kind our usual procedure was to spread a drop
of melted glycerine jelly on a slide and allow it to cool; then to dissect off the
wings (generally under water), taking with them just enough of the thorax to
include the basal attachments of the tracheae; then to place these wings upon
the solidified glycerine jelly on the slide; then to lower upon the wings a heated
cover glass, causing the jelly to melt enough to envelope the wings; and then
to cool the mount speedily on a cake of ice, a marble slab, or in a draught of cold
air. Rapid cooling is imperative, for in melted glycerine jelly the tracheze
soon become filled and the smaller ones are then invisible.’ The Amer. Nat.
Vol. XXXII, page 45.
104 Annals Entomological Society of America [Vol. II,
tion except that even the balsam mounts remained usable for a
sufficient time to secure camera lucida drawings of the points in
question. In preparations for photomicrographs the glycerine
jelly mounts were better.
After comparing the two methods the balsam mounts were
used almost altogether as they were quicker and simpler to pre-
pare and a much larger proportion of usable mounts were made
in this way.
The wings of freshly emerged aphids were severed from the
body together with a portion of the thorax, to preserve the basal
part of the tracheze and to block to some extent the penetration
of the balsam.
In some species, where the shape of the body and the angle at
which the wings are set permitted it, the aphid itself with wings
attached was mounted after beheading the insect and puncturing
the tip of the abdomen. The legs had also to be removed. The
pressure of the cover glass in these mounts generally forced the
body fluids from the openings at the two extremities and often
excellent tracheal preparations were secured in this way. It
could be applied only to a limited number of species, however.
Only wings of freshly emerged aphids are available for the
study of the wing trachez, as before the wings are hardened
enough for the first flight, the trachez have become either en-
tirely invisible or so nearly so that they are useless for the ques-
tions inhand. It thus becomes necessary to select the individuals
-in the first few minutes after molting. This can readily be done
by taking the palest insects while the wings are yet white, that is,
before they begin to become transparent. An abundance of
material was kept on hand by the collection of large colonies of
aphids whenever any were found, the wing pads of which indi-
cated that the final molt was approaching.
It was more difficult to secure the wing pads at the critical
moment. The wing pad of the aphid nymph is not only small
but it is relatively thicker and softer and narrower than those of
many insects and the developing wing is for the most part so
crumpled that the courses of the trachez are impossible to follow.
So soft is the tissue of the nymphal pad and so loose do the tra-
cheee lie within it, that the slighest pressure is likely to misplace
them or rupture them, and render the preparation useless. By
selecting the nymphs of such species as had the flatter wing pads,
1909] Homologies of Wing Veins 105
the best results were obtained. However, none of the aphids
are so easy to manipulate for the tracheation of the wing pads as
are the flat padded psyllids.
* * * * *
Although the possibilities of interesting features of wing tra-
cheation of the aphids were by no means exhausted with any
species; for the problem at issue,—the homologies of the wing
veins,—no point was left in question. Wings of more than two
thousand of newly emerged aphids were examined. Where
possible a single point was verified for all the variations of wing
types from Lachnus to Chermes. Where a point arose that cer-
tain species showed better than others it was exhaustively studied
by making numerous mounts of those species which had any bear-
ing upon it.
As for the range of material used, approximately roo species
belonging to 16 genera were drawn upon. Many of these were
discarded after a few mounts,—as for instance after the condition
for the genus Aphis was clearly ascertained other species of this
genus did not give additional data. Many species were found
unsatisfactory to work with by reason of size, density of color
or other conditions and were discarded after a few trials.
The genera from which most of the data were accumulated
and in all of which the tracheation was demonstrated repeatedly
and conclusively, are Lachnus, Melanoxanthus, Callipterus, Chai-
tophorus, Myzus, Macrosiphum, Rhopalosiphum, Aphis, Schizo-
neura, Mindarus, Pemphigus, Tetraneura, Hamamelistes, Cher-
mes and two new genera still in manuscript. These genera pre-
sent practically the whole variation of the types of aphid venation
from the more generalized to the more specialized.
The more logical presentation of the four families considered
in this paper would be in the following order, —Psyllide, Aphidide,
Aleurodidze and Coccide, but as the problem was undertaken
primarily for the Aphididz and as the investigation was devoted
for the most part to them, the part dealing with the aphids is
presented first. The work with the other families, though suffi-
cient to indicate the homologies of the veins, has been so much
slighter in amount that it seems more fitting to give secondary
place to the Psyllide even though this necessitates a break in the
systematic sequence.
106 Annals Entomological Society of America [Vol. II,
THE COURSE OF A VEIN.
ae
In the following pages the “‘course’’ and the ‘‘free part”’ of
veins are frequently mentioned. By the free part of a vein is
meant all that portion that is not coalesced with any other
vein. As for instance in Fig. 36, the free part of Cu, is all that
portion of Cu, between the point where it separates from Cu,
and the margin of the wing.
As to the course of the vein it has been convenient to consider
each of the branches of any vein as extending from the base to
the margin of the wing, as is made plain by the following quo-
tation: ds
“‘Tf radius and its five ranches be taken as an example, the
stem part, always designated as R, would be considered as being
a combination of all the branches of radius, or as R,+.,+3+,+5,
which divides into R, and R,. In like manner the stem of the
radial sector would be considered as being a combination of all
the branches of the radial sector, or as R,+3;+,+,;, which divided
into R,+, and R,+,;, and these in turn into R, and R,, and R,
and R,, respectively. So that in tracing out the course of any
of the branches of radius by drawing a pencil along them, as R,,
beginning at the base of the wing, we would pass first over the
stem of R, then over the stem of the radial sector, then over
R,+,, and finally over the free part of R,.”’
COALESCENCE OF TRACHEZ.
A comparison of the tracheation of the wing pad of an aphid
nymph with that of the wing of the recently emerged aphid of
the same species reveals the fact that there are fewer basal tra-
cheze in the later than in the earlier stage, although the terminal
branches are the same in number. For instance, there are four
tracheze which branch from the body trachea in the nymphal
wing pad of Schizoneura rileyt (fig. 21) and only two tracheal
stems at the base of the newly emerged wing of the same species
(fig. 22). Similarly as there are four tracheez at the base of the
nymphal wing pad of the species of Aphis which I have studied
and of Callipterus ulmifolit while there are but two basal tracheze
in the newly emerged wings of these same species, this is shown to
be the normal condition for the family in the more generalized
genera. In the nymphal wing pad of Mindarus (fig. 18) three
* MacGillivary, A. D., Wings of Tenthredinoidea. Proc. U.S. Nat. Museum
Vol. XXIX, page 576. 1906.
1909] Homologies of Wing Veins 107
of these trachez unite before they reach the body trachea,
approaching the condition of the mature wing.
This change in the condition of the corresponding tracheze
at different stages has been designated in this paper as the
‘‘coalescence’’ of tracheze, as this term seems to express the rela-
tion of the tracheee of the mature wing to those of the nymphal
wing pad. This sort of coalescence of tracheze is a constant fac-
tor in the aphids, not alone as to the main tracheal stems but a
similar tendency is shown in the branches of the tracheee. For
instance notice that the medial trachea in figure 21 branches
desidedly nearer the base of the wing than the medial trachea
in figure 22, these branches being coalesced to much nearer the
margin of the wing than in the earlier stage.
How or when this coalescence takes place the writer has as
yet made no attempt to ascertain. No pains has been spared,
however, in accumulating evidence that it does occur, or in mak-
ing sure that it is an actual union of trachez and not an approxt-
mation. A 1-6 objective was usually sufficient for the exami-
nation of the cases in question. Where it was not, an oil emersion
was used. The manner and time of the coalescence is a mystery
well worth solving but it is only the fact of this coalescence and
not its method which has any bearing on the present problem—
the homologies of the veins. However, it is a problem which the
writer hopes to undertake in the near future.
Besides the normal coalescence of the basal portions of medial,
cubital, and the first anal trachez just described, very rare
instances of abnormal coalescence occur. Figure 2 shows a
wing of Melanoxanthus in which the radial, medial and cubital
tracheze are anastomosed for a considerable distance.
Attention is also called to Plate XIX, which shows three unusual
examples of tracheation. Normally the forewing of Chermes
has two basal trachez as in the more generalized genera, and as is
shown in figures 24 and 28. But in this highly specialized genus
the tracheation seems to be unstable. Figure 26 shows an in-
stance of anastomosis similar to that in figure 2, while figure 27
represents a single tracheal stem at the base of the wing, a condi-
tion which approaches the normal condition of the hind wing of
Chermes (fig. 30.) On the other hand figure 25 records an
instance where the first anal trachea is separate from the common
stem of the medial and cubital trachez to the base of the wing,
giving in this case three basal trachez.
108 Annals Entomological Society of America [Vol. IT,
In the study of the tracheation in the Psyllidze the writer
has been confined to a single species, and we have here even a more
striking degree of normal coalescence of trachez than in the Aphi-
didae. For instance, as is shown in figure 33, the seven trachee.
are distinct to the body trachea in the nymphal wing pad, while
figure 34 records the normal tracheation of the freshly emerged
wing, the three most important trachez, the radial, medial, and
cubital tracheze, are coalesced into a common stem at the base,
and for a considerable distance the medial and cubital trachez.
are stalked after the radial trachea has separated from them.
We have, then, a very striking difference in the relation of the
trachez of the nymphal wing pad and those of the freshly emerged
wing, in both of these families of insects.
COSTA AND SUBCOSTA OF APHIDIDA.
The costal vein extends along the cephalic margin of the wing.
No trachea precedes this vein in the freshly emerged wing.*
Neither is a costal trachea present in any nymphal wing pad of
the species examined by the writer. Further, no evidence of a
vestigial costal spur from the body trachea is to be found in the
several species examined for this point. Perhaps the most
striking evidence of a costal vein aside from the stiffening of the
costal margin is the fact that in severing the wing from a freshly
killed aphid, the yellow body fluids frequently flow into this vein
and extend along to about the region of the stigma (fig. 4).
In like manner no trachea precedes the vein subcosta in any
species examined of the sixteen genera (see page 105) of aphids
from Lachnus to Chermes which I have studied for this point.
No item in this work has been more carefully investigated than
the possibility of a subcostal trachea.
Indications of such a trachea have by no means been lacking.
For instance, the fold of the vein subcosta appears very early in
the freshly emerged wing (long before the other veins) and is
therefore frequently the only vein indicated at the time the tra-
chee are visible. For many mounts of many species this form-
ing vein resembles a tracheal line closely enough to be mis-
taken for one unless exceedingly great care is taken to secure
* Comstock and Needham. Wings of Insects. Page 858. ‘‘* * * there
are often channels present which do not contain trachea. This is oftenest true
of two large channels at the lateral margins of the wing. Of these the costal
remains abundantly lined with cells, which ultimately form the strong costal
vein. Its trachea is often atrophied, probably owing to the disadvantageous
position of its base in relation to air supply, as we have hitherto indicated.”
1909| Homologies of Wing Veins 10g
wings immediately after emergence. Handlirsch* thus mistook
this forming vein for a trachea in his preparation of Schizoneura
lantigera.
Wherever there has been the slightest chance that a subcostal
trachea might be present, exhaustive investigation has been made
until no possible doubt of its absence remained.
Again, for some species, one of the secondary branches of the
radial trachea near its base might easily be mistaken for a sub-
costal trachea so far as its position goes. Chermes pinifolie
(fig. 26) serves as an illustration of this. But these secondary
branches are very variable even for the same species and after
examining large series of mounts little difficulty is experienced in
distinguishing the trachee of the main veins from secondary
branches.
The only trachea found during the whole time possible to
interpret as the subcosta was a single mount out of hundreds
of Mindarus (abtetinus Koch?) where an exceedingly short branch
from the radial trunk near the base occurred, larger than the
ordinary secondary trachee. This may be an abortive subcostal
not entirely atrophied. Or it may be merely an abnormality.
This species of Mindarus, collected in great abundance from Balsam fir
in the vicinity of Orono is apparently the same as abietinus Koch. But as it
has not yet been compared with specimens from Europe it is given in this paper
accompanied with a question mark.
Another opportunity for confusing a branch of the radial
trachea for a subcostal trachea is found in wings where the path
of the tracheal branch of radius chances to coincide with that of
the newly forming subcostal vein. Such a case is shown in the
tracheation of Pemphigus acerifolit (fig. 12) where the trachea
follows the stigmal margin that the subcostal vein is later to
bound. But the branches of the radial trachea are quite as likely
to cut directly across the path of the forming subcostal vein as to
coincide. Chermes abietis (fig. 24) will illustrate this.
The point, however, which gave me most trouble was the fact
that very frequently the fold in the wing caused by the forming
subcostal vein bent the branches of the radial trachea to such an
extent that their connection with the radial trachea was obscured
and a series of tracheal branches resulted which had every appear-
ance of rising directly from the region of the subcosta.
* Handlirsch, A. Fossilen Insekten. Plate VIII. Fig. 9.
TIO Annals Entomological Society of America [Vol ih,
The absence of a costal trachea is by no means unusual.* The
aphid wing is characterized by the reduction of tracheze and the
loss of the subcostal trachea is only one more step in the general
trend of reduction. What the aphid lacks in main tracheal
trunks, however, it makes up in secondary ramifications, and a
glance at the tracheation of Chermes abietis (fig. 24) will show that
the absence of costal and subcostal trachez does not indicate that
this portion of the wing is untracheated. Perhaps the abundance
of secondary trachee explains the absence of unnecessary costal
and subcostal trachee. At any rate they might in some cases be
mistaken for one of these main trachee. Refer again to Chermes
abtietis (fig. 24) where the ends of the radial branches are so
interwoven as to have the appearance of a continuous subcostal
trachea. And where the channel of the subcostal vein hides the
radial connection, we have frequently the occurrence of what
appears to be a strong subcostal trachea with branches extending
toward the radius. Further, in case the radial connection of all
these trachez, except a single one near the base of the wing, is
obscured, as sometimes happens, there is, to all superficial appear-
ances, a clear indication of a strong subcostal trachea.
Altogether the absence of the subcostal trachea was one of
the most troublesome points to establish. Each clue was fol-
lowed, however, until the evidence was absolutely conclusive.
Although no subcostal trachea is present, there seems to be
no other conclusion except to consider that the vein subcosta is
present in the large main vein channel of the wing, and extends
from the base of the wing to the stigma where it approaches the
margin of the wing. Indeed it seems not at all unlikely that this
stiff vein fold at the base of the aphid wing is responsible for the
atrophy of the subcostal trachea. The radial trachea from its
position can elude the pressure and run alongside, but this ob-
struction alone would seem sufficient to explain the loss of the
subcostal trachea in the Aphidide.
RADIUS OF APHIDIDA.
The tracheation of Myzus cerasi (fig. 3) gives a fairly typical
two-branched radial trachea. This type persists for the family
of Aphididze except in the Chermesinze where this trachea is not
branched. This trachea is one of the two main trachez which
enter the wing, and from the phylogenetic standpoint the chief
* See Footnote, page 108 (Costa and Subcosta).
1909] Homologies of Wing Veins ia
interest concerning it is centered in the cephalic branch, or the
trachea preceding vein R,.
In the case of two widely separated Hemiptera, a Cicada and
a Coreid, this trachea was found to be but weakly developed in the
nymphal wing pads,* being apparently crowded out by the strong
subcostal trachea, and it was not succeeded by a vein,—the vein
R, being completely absent in these two insects.
With the aphids no such crowding of the radial trachea occurs,
for, as has been shown, the costal and subcostal are both absent.
It is, however, of great phylogenetic interest to find that in the
tracheation of the aphids, the weak character of this same tra-
chea (that is the one preceding vein R,) is evident. In Lachnus,
probably the most generalized genus in the family, this branch is
scarcely to be distinguished from the secondary branches of the
radial trachea and is emphasized chiefly from the fact that it is
succeeded bya vein R, (fig. 1). And in all of the generalized
genera it is characterized by a wavering and uncertain course.
It is consistent with this general tendency to find that this is the
tracheal branch which if it appears at all in the specialized Cher-
mesinz is so weakly indicated as to become indistinguishable
from the secondary branches and loses its significance as it is not
succeeded by a vein, vein R, being absent in the Chermesine.
The wings of the Chermesine are at a glance conspicuously of
a different type from those of the more generalized aphids, and
this difference in venation has heretofore been interpreted as due
to the absence of media (the “third discoidal’’). Especial atten-
tion is called, therefore, to figures 28 and 29. These are two draw-
ings made of the same identical wing of Chermes pinifolie,—
figure 28 drawn immediately after mounting when most of the
tracheee showed, and figure 29 drawn from the same mount 24
hours later when the trachez had cleared and become invisible
and the veins are apparent instead. These drawings indicate
the character of the difference between the Chermes venation and
that of the more generalized genera. R, has disappeared and
Rs, losing its characteristic curve, lies along the caudal margin of
the stigma. Media, apparently to replace the position and func-
tion thus deserted by the radial sector, migrates forward to ap-
proximately the position which the ‘‘stigmal vein”’ or radial sec-
tor occupies in the generalized genera. This is an interpretation
which could in no wise be settled except by appeal to the trachea-
* Comstock and Needham. Wings of Insects. pp. 245—251.
Di2 Annals Entomological Society of America [Vol. II,
tion, and it is perhaps as pretty an illustration as can be found of
the conclusive evidence which tracheation can sometimes bear ina
doubtful case of venation.
The same condition is shown in Chermes abtetis (figs. 24and 25).
Although the radial trachea has no main branches except R,
and the sector, this trachea very frequently as is the case of the
other trachez, bears a great number of secondary ramifications.
These secondary branches are omitted in many of the drawings
accompanying this study as they are the first to clear and become
invisible,* and were frequently neglected when first attention was
concentrated on other points. However, a glance at Chermes
abietis (fig. 24) and Lachnus strobt (fig. 1) representing the genera
at the extreme ends of the Aphididz show the general condition
of these radial ramifications which are further treated under the
discussion of the subcosta: (see page 108).
With an understanding of the tracheation, little explanation
is required for the vein radius. Except for the Chermesine,
the free part of R, forms the caudal boundary of the stigma and
extends to the margin of the wing. The radial sector originates
in the vicinity of the stigma and curves strongly to near the tip
of the wing. The main stem of radius extends from the junction
of these branches to the base of the wing in a line about parallel
to the subcosta,—forming indeed a part of that strong main
composite vein channel of the wing.
In Chermesine, radius is unbranched, as has been explained,
and takes a straight course from the base of the wing to the ex-
tremity of the stigma. A double adjustment takes place here.
Correlated with the loss of R,, which in the more generalized
genera bounds the stigma along its distal edge, is a straightening
of the caudal margin of the stigma and a straightening of the .
course of the radial sector so that the radial sector runs along the
edge of the broadened stigma in this type of wing (figs. 31 and 32).
MEDIA OF APHIDIDA.
For a correct interpretation of media an understanding of the
tracheation is necessary. The freshly emerged wing of Myzus
cerast (fig. 3) gives a fairly typical tracheation for the more gen-
eralized wings. The medial trachea will here be seen, as'is true
for all the genera of Aphididz, to be the second of the two main
tracheee to enter the wing from the body cavity. This trachea
* See discussion of. preparation, page 103.
1909] * Homologies of Wing Veins 113
lies uniformly parallel to the radial trachea, usually separated
from it by an appreciable distance, though often touching it, but
not, except in rare instances,* really coalesced with the radial
trachea. As we approach the wings most specialized by reduc-
tion of tracheze, we come first to Schizoneura (fig. 22) and Min-
darus (fig. 19) as examples of the medial trachea with but two
branches, and then to such wings as those of Pemphigus, Tetra-
neura, Hamamelistes and Chermes, where media is unbranched.
The migration of the medial trachea to approximately the position
occupied by the radial sector which in turn migrates to meet the
margin of the stigma, is characteristic of the Chermesinz, and has
just been discussed under the treatment of radius.
The tracheation of the adult wing, however, does not suffice
to make perfectly evident the relation of media to cubitus. For
instance, so far as has yet been demonstrated, what is here inter-
preted as M,+, (fig. 22) might perhaps be understood as cubitus
coalesced with media as Vickeryf has explained it, or what is
here interpreted as cubitus might be homologized as a branch of
media as Handlirscht has done. Fortunately the tracheation of
the wing pad of the nymphs of certain species make this point
entirely clear. Reference to the nymphal pad of Schizoneura
rileyt (fig. 21) gives an unmistakale demonstration concerning the
point at issue. Four main trachez are seen to be separate to the
place of connection with the body trachea. These are the fore-
runners of radius, media, cubitus and the first anal vein. The
two-branched media and the unbranched cubitus of Schizoneura
are thus clearly seen to be preceded by distinct trachez separate
to the base of the wing pad, and the relation of media to cubitus
in the mature wing becomes at once apparent and the degree of
basal coalescence evident.
After the foregoing discussion of the tracheation little remains
which needs to be said concerning the vein media. For approxi-
mately one-half the length of radius media is coalesced with
R+Sc.|] and nearer the base of the wing similarly with the proxi-
* For discussion of coalescence and anastomosis of the trachez, see page 106.
+ Vickery, ’08, page 9. Figure 2.
t Handlirsch. Die Fossilen Insekten und die Phylogenie der Rezenten For-
men. 1906-08. Taf. VIII, figures 9 and 10.
|| ‘‘The figures show that in some cases what appears as a single vein is
formed about two closely parallel trachee. This is shown in the case of the
bases of the second and third principal trachee, counting from the costal margin
of the wing, the radial and medial trachee. This illustrates a fact of frequent
occurrence, that what appears to be a single vein may be formed by the coales-
-cence of two primitive veins.’’ Comstock—Needham, Wings of Insects, p. 48.
114 Annals Entomological Society of America [Vol. IT,
mal portions of cubitus and the first anal vein. The free part of
media occupies about the center of the distal half of the wing.
Its maximum number of branches is three, —M, and M, and
M,+, (fig. 8). The first step at reduction is in such wings as
Schizoneura and Mindarus where M, and M, are coalesced, giving
a two-branched media M,+, and M,+, (figs. 20 and 23). These
two branches in turn are coalesced, resulting in the unbranched
media (M,+.+M,+,)* in the wings having. the most reduced
venation.
The connection between the free parts of media, cubitus and
the anal vein and the composite main vein is frequently broken,
on account, probably, of the strong ridge in the wing caused by
this vein. This part of media being in the central portion of the
wing is not, apparently, so necessary to the strength of the wing
as the veins on either side, and starting from the broken basal
connection media is found in various degrees of atrophy, in some
species one-third to one-half of the base of the free part of media
being lacking (fig. 17).
CUBITUS OF APHIDIDA.
Cubitus is present in all the genera of Aphidide and in all of
them unbranched.
The tracheation of the freshly emerged wing and of the wing
pad explains cubitus and its relation to the other veins.
Attention is called to figure 22, Schizoneura rileyi, which gives
a fair illustration of the cubital trachea. Its basal portion is co-
alesced with those of the medial and anal trachee. In most
wings the free portion of the cubital trachea originates about mid-
way between the medial and first anal tracheze. In some instances
however, the cubital trachea runs along parallel with the medial
trachea for some little distance. Figure 9, a mount of a newly
emerged hind wing of Macrostphum ptst, is an unusual instance
even for the species in question, where the cubital trachea runs
parallel with the medial to almost the base of the wing before it
becomes coalesced. On the other hand, in the wing of Hamame-
listes spinosus (fig. 15) a greater degree of coalescence has taken |
place and the cubital and first anal tracheze branch from the me-
dial on a common stem, and proceed some little distance before
separating.
* See page 106 for a discussion of the course of a vein.
1909] Homologies of Wing Veins iS
The tracheation of the wing itself, however, is not sufficient
to define cubitus beyond question, and numerous mounts of
nymphal wing-pads were examined with this end in view. Per-
haps the most satisfactory is that.of Schizoneura rileyi (fig. 21)
in which a portion of the body trachea is seen with four main
wing trachee, the fore-runners of radius, media, cubitus and the
first anal vein.
Cubitus is here clearly shown in its relation to media and the
first anal and the peculiar complications of the venation of the
adult wing are thus explanable as coalescences with media on the
one hand or the first anal on the other. Schizoneura rileyi was
the only species in which the writer has succeeded in getting the
connection of these four nymphal wing trachee with the body
trachea, but mounts of Callipterus ulmifolit showed four main
tracheze distinct to the base of the pad, as did also mounts of
Aphis sp. In the last nymphal stage there is a tendency, in
some species at least, for the basal portions of the medial, cubital
and first anal tracheze to become coalesced, giving the two main
tracheal stems of the mature wing. This is shown in a wing
pad of the last nymphal stage of Mindarus (fig. 18) where the
four main trachee are distinct nearly to the base of the pad where
three of them become coalesced before reaching the body trachea.
The vein cubitus so closely follows the cubital trachea that
a further discussion concerning it is not necessary.
THE ANALS OF APHIDIDA.
The same mounts which explain cubitus show just as clearly
the first anal and its relation to the other veins. We have then
the unmistakable homologies of four of the wing veins of Aphidi-
dee traced in the courses of the trachez of the freshly emerged
wings and the wing pads of the nymphs: the ‘‘first discoidal’”’
being the first anal, the ‘‘second discoidal’’ being the cubitus,
the “‘third discoidal” or “‘cubital vein’’ being the media and the
‘fourth discoidal”’ or “‘stigmal vein’”’ being the radial sector.
This seems enough to ask of the trachez of a highly specialized
wing, but they do tell even more. They give constantly the sec-
ond anal which appears in the freshly emerged wings for all the
genera studied by the writer, as a delicately marked and fairly
regularly placed trachea. No vein is formed about this trachea
so that it has no other significance for the venation than to help
determine that the anal vein here homologized as the first anal
116 Annals Entomological Society of America [Vol. IT,
has not been so homologized without tracheal evidence that it is
not the second or third anal,—a statement further elucidated by
the appearance of a third anal trachea in many freshly emerged
wings. Figures 1 and 3 and others give the second anal trachea
in its characteristic position, and figures 2, 4, 6 and others give
a few of many mounts showing the third anal trachea also. These
last anal tracheze are among the first of the tracheze to become
invisible, so that in drawings made with other points in view,
they are frequently not indicated, as they had cleared before the
other points at issue had been finished.
The presence of the second and third anal trachez in the nym-
phal wing pad has not been established in those species of aphids
in which the nymphs were studied. What the history of these
trachee is, then, previous to their appearance in the recently
emerged wing is not yet ascertained.
THE HIND WING OF APHIDIDA.
The hind wing of Chaitophorus populicola (fig. 7) will serve as
an illustration of the tracheation of the hind wings of the more
generalized genera. In figure 9 of the Macrostphum pis the four
trachez-are distinct to very near the base of the wing. Usually
a greater degree of coalescence has taken place even for this
species.
By referring this wing to the front wing of Chermes (Plate XIX)
we can readily homologize the unbranched radial trachea, the
simple medial trachea, the cubital trachea and the, first anal.
Like the second anal of the fore wing, the first anal of the hind
Wing appears only as a trachea and is not followed by a vein.
The same trachee occur in the other more generalized genera,
and also in Pemphigus (fig. 13) where the front wing is specialized
by the reduction of veins in media. When we reach Chermes
(fig. 30), however, we find, in spite of the wealth of secondary
tracheee, only the radial and medial trachezee marking the courses
of succeeding veins and these are coalesced at the base.
The venation of the hind wing is, then, homologized, as follows:
the marginal costa and the very faint subcosta, both of which,
due to the lack of mechanical necessity found in the front wing, are
not well developed; the radius represented by the radial sector
only; the media always simple; and in the more generalized
genera, cubitus. The venation of Tetraneura (fig. 17) is of
interest. The front wing has the venation of Pemphigus and
1909] © Homologies of Wing Veins Fry
the hind wing lacks the cubitus and is, therefore, a step in the
direction of Chermes.
Although we find the same veins in the hind wings of Macrosi-
phum solanifolit (fig. 8) and Pemphigus acerifolie (fig. 13) for
instance, these two wings appear different on account of the
difference of position of the veins. The radial sector takes the
same course in both wings, but the free part of cubitus originates
much nearer the base of the wing in Macrosiphum while usually
in Pemphigus cubitus is coalesced with media to the point where
media branches off from the course of radius. This peculiarity
is correlated with a similar manifestation in the front wing of
Pemphigus and is doubtless due to the long narrow wings of this
genus. |
In Chermes (figs. 30 and 31) as was stated in the discussion of
the tracheation, only veins radial sector and media are repre-
sented, and the hind wing of Phylloxera (fig. 32) has retained only
the radial sector.
PSYLLIDA.
The tracheation of the psyllid Wing is instructive both because
it throws light upon the interpretation of the venation of the
psyllid wing and because it bears strong additional evidence in
support of the conclusions arrived at for the aphid wings.
Fig. 33 gives the tracheation of the fore-wing pad of the nymph
of Psylla floccosa Patch. It will be seen by reference to this that
there is no coalescence for these trachee at this stage. They all
branch separately from the body trachea. There are seven tra-
chee: the costal; a short trachea lying at the very margin of the
wing pad; the unbranched subcostal extending in a line about
parallel to the costal margin of the pad and lying very close to
the costal trachea at the base; the radial, terminating in two long
branches, the forerunners of R, and Rs; the two-branched medial:
the typical two-branched cubital; and two anal trachee.
This comes surprisingly close, for a wing so highly specialized,
to the hypothetical type of Comstock-Needham.
The newly emerged wing shows a decided change in the appear-
ance of the trachez, but the change is due to basal coalescence
instead of any radical change in the number or branching of the
trachee.
The tracheation of the freshly emerged wing of Psylla floccosa
(Fig. 34) shows the degree of this change. The costal trachea has
118 Annals Entomological Society of America [Vole hr
disappeared as is frequently the case.* The subcostal and sec-
ond anal trachee have migrated to very near their respective
margins of the wing, but are both distinct. The first anal, a
slender but distinct trachea, follows the course of the claval su-
ture. The proximal portions of the radial, medial and cubital
trachee are coalesced into a common stem at the basal fourth of
the wing. This main stem divides into two trachez, the radial
and the common stem of the medial and cubital. The latter soon
separates again into the free portions of the medial and cubital
trachez.
There is a greater degree of coalescence in the tracheation of
this psyllid:- wing than occurs in that of the aphid wing as the
radial trachea is always (except in very unusual variations as
cited, page 107) separate to the base of the freshly emerged aphid
wing.
The venation of Psylla floccosa (fig. 36) coincides with the
tracheation of the freshly emerged wing very closely. Subcosta
strengthens with the costa the cephalic margin of the wing. The
second anal in a similar way reinforces the anal margin at the
base. The first anal is so delicately traced along the claval suture
that it cannot function as a strengthening rod in the wing as do
the other veins. Indeed it has frequently been considered not a
true vein.
Radius, media, and cubitus share about equally the burden of
strengthening the wing. They are coalesced as a common main
stem at the base. The relation of media and cubitus is similar
to that of the same veins in the wing of the aphids and this typi-
cal two branched cubitus bears further evidence of the correctness
of the interpretation of cubitus in the aphid wing.
R, responds to the mechanical necessity for strengthening the
front of the wing. It is the only vein present in this region and
upon it falls the burden as well as the approximate position usually
assumed by the subcosta in many insect wings. Ks is a strong
vein and reaches the margin near the apex of the wing.
The tracheation of the freshly emerged hind wing of Psylla
floccosa resembles that of the front wing closely. (Fig. 35).
The subcostal and second anal trachez lie near their respective
wing margins, as in the front wing. The first anal trachea occu-
* “Tts trachea is often atrophied, probably owing to the disadvantageous
position of its base in relation to air supply, as we have hitherto indicated.”’
Comstock—Needham: Wings of Insects, page 858.
1909] Homologies of Wing Veins I19
pies the same position as in the front wing. Similarly there is a
common stem of the radial, medial and cubital trachee. The
medial trachea, however, is unbranched, and the tracheal branch
preceding R, is wanting.
The loss of R, is significant. This vein is one of the least
stable of the Hemipterous wing veins. Its complete absence has
been established for the Cicada and a Coreid.* In the front wing
of the aphids R, was seen to be preceded by a wavering
trachea in all the subfamilies except the Chermesinz where the
vein R, is altogether lacking. In the hind wing of none of the
aphids does either the vein R, occur, or the corresponding trachea.
In the front wing of the psyllid and Aleurodicus alone for this
group is R, a strong vein and here it is evidently the responsé to
the mechanical necessity not otherwise provided for. Subcosta.
is wanting and, as the only vein present in this portion of the
wing, the burden of support falls upon the radius.
It is exceedingly interesting to find the condition of R, as
predicted for the Hemipterat on the basis of the Cicada and
Coreid, fulfilled in this group of highly specialized Homoptera.
It is interesting, too, to find the same veins, vein for vein,
appear in the wing of the psyllid that we have in the wing of the
aphid—the most striking difference is a very slight one,—the
cubitus branched in the psyllid and unbranched in the aphid.
There is experienced no difficulty, as has been seen, in homologiz-
ing either the aphid or the psyllid veins independently of each
other but the fact that in doing so the conclusions arrived at with
each support and bring additional evidence for the other should
in no wise be disregarded.
The fact that the costal margin of the psyllid wing is strength-
ened in one of two distinct ways (1) by a stigma and (2) by R,, is
very well brought out in the five psyilids which have been selected
to represent the venation of this family. Of these Euphalerus
nidifex Schwarzt (fig. 38) would seem to be the most generalized
type so far as the radial region is concerned, R, in this wing extend-
ing to near the tip of the wing as the trachea preceding R, does
in the nymphal wing pad of Psylla floccosa (fig. 33). In Pachy-
* Comstock and Needham: Wings of Insects. Page 245.
{+ Comstock and Needham: Wings of Insects. Page 245.
¢ For an opportunity to study and figure the wings of E. nidifex and A. mori,
the writer is indebted to Dr. L. O. Howard, who kindly loaned specimens of
these species for this purpose.
120 Annals Entomological Society of America [Vol. II,
psylla c. mamma Riley (fig. 39) the tip of R, has migrated slightly
toward the middle of the costal margin. In both these wings the
space from the base of the wing to the tip of R, is evidently too
long to bear the strain of flight without further strengthening.
This needed reinforcement occurs in the presence of a more or
less pronounced stigma the proximal edge of which is margined
by a ridge which is in some species so clearly defined as to be
frequently figured as a vein.
As the tip of R, approaches the middle of the costal margin
as in Anomoneura mort Schwarz (fig. 40) and even more in Psylla
floccosa (fig. 36) the need of the stigma is removed as R, strength-
ens this portion of the wing margin.
The wing of Trioza (fig. 37) shows an extreme case of the mi-
gration of R,, this vein being here scarcely longer than the stig-
mal ridge in Pachypsylla and in about the same position. Cor-
related with this condition are other striking-departures from the
more generalized psyllid wing, perhaps the most conspicuous being
the origin of the free parts of media and cubitus at approxi-
mately the same point as the origin of the free part of radius,—
the common stem M+Cu present in most psyllids being lacking
in Trioza.
The wing of A. mort is in one respect most unusual for the
family, and that is in the branching of the radial sector. This
is the only instance in the four families discussed in this paper
where the radial sector is branched, and it throws additional
evidence on the interpretation of this vein as the sector. It
should be stated in connection with the discussion of figure 4o,
that the venation here shown may possibly be not normal for
this species. The writer had access to but a single specimen
and in this the radial sector of the wing on one side bore five
branches as figured while the radial sector of the other side was
six branched.
The purpose of this study in wing tracheation has had the
vein homologies as a goal and is not intended to enter into syste-
matic discussions. However, since the systematists of the
Psyllidee build their tables largely upon the basis of the wing
veins,* it is apparent that it would be more satisfactory to apply
* Low. Verh. g—b. Wien XXVIII.
Maskell: Transactions of the N. Z. Institute Vol. XXII, 1889, page 158.
Frogatt: Australian Insects.
Kuwayama: Trans. of the Sapporo Nat. Hist. Soc. 1907-08.
1909] Homologies of Wing Veins Tag
a terminology which has the same significance for not only the
closely related families of Aphididae, Aleurodide, and Coccide,
but the other insect orders as well.
For instance it would certainly seem more convenient to say
‘‘M-+Cu is longer than the stem of R”’ than to resort to the more
tortuous statement of Maskell*: ‘‘the stalk of the lower branch
(cubitus) of the furcation of the primary vein is longer than the
stalk of the upper branch (subcosta).”’
At present the veins in the figures of the psyllid wings are
indicated by arbitrary letters or figures with no necessary relation
to symbols used by the same author for any other psyllid wing
or to those of any other author for the same wing. This hap-
hazard arrangement of lettering or numbering the figures of
psyllid wings increases the confusion caused by the fact that
the veins themselves are cumbered with such a system of nomen-
clature as the following:
“Stalk of cubitus, lower branch of cubitus, lower fork of lower
branch of cubitus, upper fork of lower branch of cubitus,”’ etc.
But by the use of a uniform system of wing terminology the
abbreviation of the names of the veins become the natural and
inevitable symbols to use for lettering the figures of the wings,
and no confusion arises in associating Rs of the figure, for instance,
with the radial-sector of the text.
The relative simplicity, ease of abbreviation and uniformity
of such a system of terminology is recommendation enough aside
from the homological significance it bears.
As is stated in the discussion of Redtenbacher’s homologies
(see page 125) except for his interpretation of the alternate con-
cave and convex veins his terms for the psyllid veins are in the
main those applied in this present paper upon the basis of the
tracheation.
ALEURODIDA.
Four fine but distinct trachez are present in the freshly
emerged wing of Aleurodes sp.,—the costal, subcostal, radial and
cubital trachez (fig. 44). All of these are uncoalesced to the
* Maskell: Trans of the N. Z. Institute, Vol. XXII, 1899, page 158. ;
{ The Comstock-Needham system of terminology has been adopted by
Handlirsch in all his recent papers dealing with venation of fossil insects and in
his extended monograph of the fossil insects of the world. (Die Fossillen Insek-
ten und die Phylogenie der Rezenten Formen 1906-1908). It has been with
interest that the writer has noticed the application of this system of nomenclature
to the group of insects with which this present paper deals, especially as the basis
for his conclusion was a study of the treacheation of the wings. ~
122 Annals Entomological Society of America [Voli |
base of the wing. The medial trachea is suggested merely by a
very faint and delicate but constantly appearing tracing in the
wing.
As in the wings of Aphididae and Psyllide the radial trachea
in Aleurodide is branched, being represented by the branches
corresponding to R, and Rs. And as in the hind wing of all
aphids and the front wing of the Chermesinz, and the hind
wing of psyllids, the vein radius in Aleurodes is unbranched, R,
‘being lacking. The formation of the vein radius in Aleurodes is
of exceeding interest. It follows the course of the radial trachea
.to the branching of the trachea and then proceeds along the radial
sector. R,in the mature wing is lost and its position, if indicated
at all, is suggested by the most delicate ‘“‘shadowing”’ in the wing
tissue. As has been previously stated,* the ‘“‘complete absence
of vein R,” was predicted as characteristic of the Hemiptera on
the basis of the two widely separated insects,—the Cicada and a
Coreid, and the phylogenetic significance of the added testimony
of the weakness of this vein and the trachea that precedes it,
in the remote and highly specialized group of Homoptera here
under consideration seems to the highest degree interesting.
The tracheation of Aleurodicus I have not had.an opportunity
‘to study. On the basis of figures of the mature wing (not a safe
basis for this group by any means, as has been shown), however,
the additional vein found there can only be interpreted as R,.
Its prominence is to be accounted for here doubtless, as in the
case of its prominence in the forewing of the psyllid, by the round-
ed shape of the wing which results in a large wing expanse not
otherwise strengthened. In response to mechanical necessity
this vein, weak or absent when not needed, becomes more strongly
developed. « [a
The second vein of the wing of Aleurodes is cubitus as will be
seen by comparing figures 44 and 45 where the second vein is
shown to follow the course of the cubital trachea.
The hind wing of Aleurodes (fig. 46) has but one trachea and
one vein,—the radial sector it seems inevitable to conclude.
COCCIDA.
Before examining wings of freshly emerged male coccids it
seemed possible only to echo the sentiment of Redtenbacher,—
‘‘Cocciden konnte ich nicht untersuchen.’’} Imagine the surprise,
meee) rage 119:
+ Vergleichende Studien uber das Flugelgeader der Insekten. Page 188.
. T909] Homologies of Wing Veins 523
then, which was caused when mount after mount of Dactylopius
Sp. on cactus showed five very delicate but perfectly distinct
trachez. See Figure 42.
Four of these trachez are simple and uncoalesced to the base
of the wing. They are arranged in two groups, the base of the
subcostal, radial and medial trachez lying close together and the
cubital and first anal tracheze forming the other group at a little
distance from the first.
The subcostal and radial trachez are both very wavy and as
they le close together, they cross and recross, often for the
greater part of their length.
The medial trachea takes a median course. This consists of
two interwoven branches, pursuing a common course.
The trachez in the wing of this coccid remain distinct until
after the veins begin to form so that the relation of the two is at
once discerned.. One vein follows the general trail of the sub-
costal and radial tracheze. This vein very evidently represents
radius.
The second vein follows the base of the first tracheal group to
about the point where the medial tracheal separates from the
subcostal and radial. The vein here takes a direct line for the
middle of the caudal margin of the wing. For slightly less than
one-third the length of this vein it frequently joins the path of the
cubital trachea. This corresponds most closely with media.
Besides these two main veins a short spur representing the
subcosta is present.
In a wing so highly eNeeranae as the coccid wing it is not
improbable that the tracheation has lost its value as a basis for
the venation. Certainly in the species studied there seems no
necessary connection between the tracheze and the veins which are
found later.
In Pseudococcus citrt the tracheation was exceedingly diffi-
cult to trace. The same veins occur in this species (fig. 41) as
in the preceding (fig 43).
In many species of Coccids there are shadowed portions of the
wings. It is due to this fact that we find the coccids sometimes
figured with apparently four long veins, alternately dark and light,
as for instance, in Westwood.*
* Westwood, J.O. Arcana Entomologica. Vol. I, Plate 6.
124 Annals Entomological Society of America [Vitals
HISTORICAL DISCUSSION.*
As is the case with other groups of insects the names applied
to the wing veins of aphids were given in the first place without
homological reference to the veins of other insects. That is,
we have an arbitrary system which has no significance outside
the family of Aphidide. Recently several papers have appeared
which rename the wing veins with this aim in view, but as the
true homologies of the wing veins themselves have not heretofore
been established, certain mistakes in applying the Comstock-
Needham nomenclature have, of necessity, arisen. These papers
will be discussed in turn. First, however, it would seem of inter-
est to review the earlier terminology.
For convenience in this discussion attention is called to Fig.
8, in which the names are used as based upon the homologies re-
vealed in the study of the tracheation of the wings as recorded
in this paper. |
The following table will show the relation of the Comstock-
Needham terminology to the terminology previously in use for
the wing veins of aphids:
After Buckton Current terminology Comstock-Needham
Fore Wing (See fig. 8)
Costal Nervure Costal Costa (C.)
Cubitus or post-cos- Subcostal vein Subcosta (Sc)
tal nervure Radius, (R,) (together
with basal portions
of the remaining wing
veins).
Stigmatic Stigmal vein Radial sector (Rs)
First furcal »”. First branch Media (M,)
Second furcal Second branch Media (M.,)
Cubital nervure Third discoidal or Media (M)
cubital
Media ,+, (M,+,)
Second oblique Second discoidal Cubitus (Cu)
First oblique First discoidal First Anal (1st A.)
Hind Wing.
Cubitus or post Subcostal vein Radial sector (Rs).
costal nervure
Second oblique Second discoidal Media (M.)
First oblique First discoidal Cubitus (Cu.)
* For careful general historical discussion of studies of wing-venation see
MacGillivrary, A. D. (’06). Wings of the Tenthredinoidea. PP. 570-574.
1909] Homologies of Wing Veins 12
Fat
The interpretation of the so-called subcostal has been*
that it is a single heavy vein spreading at the distal portion into:
the stigma. That this interpretation is incorrect the foregoing
discussion on the tracheation has made clear. [It is really a
composite structure bordered along the cephalic margin by
the subcostal vein and containing the radius and the coalesced
proximal portions of the remaining wing veins.
It will be seen by reference to the accompanying table that
the first, second and third discoidals are respectively the 1st Anal,
Cubitus and Media. The second anal not persisting in the adult
wing (that is, it is represented only by a constantly appearing tra-
chea which soon clears) is not shown in any published figures of
aphid wings.
In the hind wings the first and second discoidals are respective-
ly (fig. 8) the cubitus and media, and the so-called subcostal vein
is the radial sector.
Redtenbachert gives homologies of the wing veins of aphids
and psyllids, which except for his interpretation of alternate
convex and concave veins, come pretty close to the conclusions
arrived at from the study of the tracheation. Except for Sc and
vein VI in the forewing and that Sc is not accounted for in the
hind wing his psyllid wings are correct. His difficulties with the
aphid wings could not be settled without reference to the tra-
cheation.
Handlirschf} gives the only figure of wing tracheee for psyllids
or aphids published previous to this present paper. His inter-
pretation of the trachee for the wing pad of the psyllid nymph
is practically the same as that given in this paper. With the
freshly emerged wing of Schizoneura langigera, however, three mis-
takes occur. The line which he indicates as the subcostal tra-
chea does not occur as a trachea but as a well defined fold of the
subcostal vein. (See page 108). The unbranched trachea which
he considers M is really Cu. and his Cu. is one of the anals. The
question in regard to M. and Cu. however could only be settled
by the testimony of the tracheation of the wing pad. (See page
fi3 and fig. on):
* Oestlund, O. W. Aphididae of Minnesota, p. 4. 1887.
Comstock, J. H. Introduction to Entomology, pp. 158-159. 1888.
+ Redtenbacher, Josef. Ann. k. k. Naturh. Hofmus., I. 1886.
t Handlirsch, Anton: Die Fossilen Insekten und die Phylogenie der Rezen-
ten Formen. 1906-08.
126 Annals Entomological Society of America [Vol ie
Handlirsch homologizes the veins of the Phytophthires which
he figures and also names what trachez he figures.
In Woodworth’s discussion of the wing veins of Phytoph-
thires* the veins of these insects are homologized to the extent
that the statement is made that the front wing of Aleurodidz
is the same 1n its Maximum condition of venation as the maximum
of the hind wings of Aphidz, and the hind wings the same as the
minimum of Aphid, and that the venation of the front wing of
Coccide is the same as the minimum of the front wing of Aleuro-
dide. This author figures wings of all the. Phytophthires he
discusses but as the wing veins are not labeled and it is not appar-
ent from the text which of the ‘‘independents”’ he interprets
the media and the cubitus to be either with the. psyllids or
aphids it is not possible to homologize them from his figures.
The meaning of his statement that in only the Psyllide is the
venation extensive, enough to show clearly their affinity
to the higher Homoptera is not apparent, as his maximum
number of veins for the aphids according to his figures exceeds
the maximum for the psyllids; and: the number when the mini-
mum is taken is the same for both wings except for one extra
branch for the psyllids. Exception, moreover should be taken to
his diagram of the venation of the Aphidide, for the ‘‘additional
independent and the cross vein at the tip’’ which he states are
very rarely present, are never present in fact in the normal vena-
tion of any species of aphid. And if abnormal or freak veins are
to be included in the venation of aphid wings there would be no
reason to stop at these two, for cubitus might be branched, any
of the branches of media have an extra fork and ‘‘cross-veins”’
might be almost indiscriminately located as between cubitus
and the first anal.
In a very convenient classification of some of the external
characters of Aphidide Mr. Vickery} attempts a homology of
the wing veins of aphids with those of other insects, stating that
the names he gives, based upon Comstock’s system, were decided
upon from a study of other reduced wings such as Psocus, and
from a study of the abnormal venations found so frequently in
plant-lice. Mr. Vickery’s homologies are based upon the com-
parison of mature wings and the misinterpretations which have’
* Woodworth, C. W. (06). The Wing Veins of Insects. pp. 124-125.
. ¢ Vickery, R. A. ('08). A Comparative Study of the External Anatomy
of Plant-lice. .
T909| Homologies of Wing Veins me
“I
arisen are readily explained by this fact. Perhaps in no other
family could be found a clearer demonstration of the value of the
study of the tracheation in this connection.
SUMMARY.
The introductory discussion outlines the reasons for under-
taking this study of wing vein homologies, states the conditions
under which the study has taken place, and mentions some of the
difficulties involved in the manipulation of this sort of material.
Attention is called to the coalescence of trachea both the nor-
mal occurrence and some unusual instances.
All the veins present in the four families discussed are homo-
logized, each being taken up for detailed consideration and the
tracheal basis for the homologies being carefully stated.
The historical discussion includes a notice of all such published
work as has any direct bearing upon this present paper.
All the main features of venation or tracheation discussed are
illustrated by the accompanying figures.
BIBLIOGRAPHY.
Buckton, G. = Monograph of British Aphides.
Comstock, J.H. Introduction to Entomology.
Comstock, J. H. Manual.
Comstock, J. H. and Needham, J. G. Wings of Insects.
Froggatt, W.W. Australian Insects.
Handlirsch, A. Die Fossilen Insekten und die Phylogenie der Rezenten Formen
1906-1908.
Kuwayama,Ivon S. Die Psylliden Japans. Trans. of Sappro Natural Hist.
Society. 1907-08.
Loew, es Gen. Aphalara et Rhenocola. 1882. (Zoologisch Botanische Wien
AetheeXc I.)
Loew, F. Mittheilungen tiber Psylloden. Verh. z—b. Wien XXIX, pp. 549-
598. pl. XV. 1879.
Loew, F. Psylloden. Taf. 1X. (1878.) Verh. z—b. Wien XXVIII.
Loew, F. Revision der palaarktischen Psylloden in Hinsicht auf Systematik und
Synonymie. Verh. z—b. Wien XXXII, pp. 225-254. 1882.
Loew, F. Zur Biologie und Charakteristik der Psylloden nebst Beschreibung
zweier neuer Species der Gattung Psylla. Verh. z—b. Wien, XXVI, pp.
187-216. pls. 1 and 2, 1876.
Loew, F. Zur Sy stematik der Psylloden. Verh. z.—b. Wien, XXVIII, pp. 586—
610, pl. IX. (Figures of the neuration of 11 genera) 1878
MacGillivray, A.D. <A Study of the Wings of the Tenthredinoidea. Proceed-
ings of the Nat. Museum. Vol. XXIX. 1906.
Oestlund, O. W. Aphididz of Minnesota. 1887.
Packard, A.S. 5th Report Ent. Commission.
Redtenbacher, J. Flugelgeader der Insecten. Ann. k. k. Naturh. Hofmus. I.
1886.
Riley, C. V. Noteson N. A. Psyllide. Biological Society of Washington, 1883.
Sanborn, C. E. Kansas Aphidide.
Schwarz, E. A. Entomological Soc. of Washington. 1902-04. Notes on
North American Psyllide.
Scott, J. Monograph of the British species belonging to the Hemiptera~-Homop-
tera, family Psyllide; Tr. E. Soc. 1876. pp. 525-569, pls. VIII and IX.
128 Annals Entomological Society of America [Vol. IT,
Scott, J. On certain Genera and Species of the Group of Psyllidz in the col-
lection of the British Museum. 1882. Trans. Ent. Soc. London. 1882.
pp. 449-473. Plates XVIII, XIX.
Scudder, S. H. Fossil Insects.
Vickery, R. A. A Comparative Study of the External Anatomy of Plant Lice.
1908.
Woodworth, C. W. The Wing Veins of Insects. 1906.
EXPLANATION OF PLATES.
The figures submitted have been selected from numerous ones prepared
in the progress of this study. In their preparation two methods were employed.
The figures showing tracheation are from camera lucida sketches verified or
corrected by examination of each significant point through a high power lens,—
oil emersion when necessary. The figures giving venation (except Fig. 29) are
ink tracings of photographs,—the blue prints which were used being bleached
after the drawings were made in order that black and white figures uniform with
the camera lucida sketches might be secured.
PLATE XVI.
Fic. 1. Lachnus strobi (Fitch). Tracheation of wing of newly emerged
aphid. See page 111.
Fic. 2. Melanoxanthus sp. Wing of newly emerged individual showing
an unusual case of coalescence of trachee, See page 107.
Fic. 3. Myzus cerasi (Fab.) Tracheation of wing of newly emerged
aphid. See page 110.
Fic. 4. Aphis species. Tracheation of wing of newly emerged aphid
showing connection with the body trachea. The region of the costal vein and
also the main vein channel of the wing indicated here. See page 108.
Fic. 5. Callipterus ulmifolii Monell. Tracheation of wing of newly emerged
aphid.
‘i Fic. 6. Chaitophorus nigre Oestlund. Tracheation of wing of newly
emerged aphid. See page 116.
Fic. 7. Chaitophorus populicola Thos. Tracheation of hind wing of newly
emerged aphid. Typical, showing connection with body trachea. See page 116.
Fic. 8. Macrosiphum solanijolii Ashmead. Venation of both wings.
Fic. 9. Macrosiphum pisi. Tracheation of hind wing of newly emerged
aphid. Very exceptional. See page 116.
PLATE XVII.
Fic. 10. Microparsus variabilis Patch. Venation of both wings. Vena-
tion of this species very unstable. The type figured is a common one.
Fic. 11. Toxoptera graminum Rond. Venation of both wings.
Fic. 12 and 13. Pemphigus acerifolii Riley. Tracheation of fore and hind
wings of newly emerged aphid. See pages 109 and 117.
Fic. 14. Pemphigus venafuscus Patch. Venation of both wings.
Fic. 15. Hamamelistes spinosus Shimer. Tracheation of wing of newly
emerged aphid. See page 114.
Fic. 16. Hamamelistes spinosus Shimer. Venation of both wings.
Fic. 17. Tetraneura sp. Venation of both wings.
PLATE XVIII.
Fic. 18. Mindarus (abietinus Koch ?). Tracheation of wing pad of
nymph. See page 115.
Fic. 19. Mindarus (abietinus Koch ?). Tracheation of freshly emerged
wing.
Fic. 20. Mindarus (abietinus Koch ?). Venation of both wings. See
page 114.
Fic. 21. Schizoneura rileyi Thomas. Tracheation of wing pad of nymph.
See page 114.
Fic. 22. Schizoneura rileyi Thomas. Tracheation of freshly emerged
wing. See page 114.
Fic. 23. Schizoneura americana Riley. Venation of both wings.
T909| ' Homologies of Wing Veins 129
PLATE XIX.
Fic. 24. Chermes abtetis (Linn.). Tracheation. Note how the branches
of the radial trachea weave along the margin. See page 110.
Fic. 25. Chermes abietts (Linn.). Tracheation. An extraordinary case
in which the first anal trachea is distinct to the base of the wing. See page 107.
Fic. 26. Chermes pintfolte Fitch. Tracheation. An unusual case of
coalescence. See page 107.
Fic. 27. Chermes pinifolie Fitch. Tracheation. An unusual case of
coalescence. See page 107.
Fic. 28 and Fic. 29. Chermes pinifolie Fitch. Two camera lucida draw-
ings of the same identical wing. Fig. 28 shows the tracheation of the freshly
emerged wing, and Fig. 29 shows the course of the veins Rs, M, Cu and Ist A,
twenty-four hours after Fig. 28 was drawn and after the trachee had all become
invisible. See page 111.
Fic. 30. Chermes pintfolie Fitch. Normal tracheation of hind wing. See
page 116.
Fic. 31. Chermes. Venation of both wings.
PLATE XX.
Fic. 32. Phylloxera sp. Venation of both wings.
Fic. 33. Psylla floccosa Patch. Tracheation of wing pad of nymph.
Fic. 34and Fic. 35. Psylla floccosa Patch. Tracheation of freshly emerged
fore and hind wings.
Fic. 36. Psylla floccosa Patch. Venation of fore wing.
Fic. 37. Triozga species. Venation of both wings.
PLATE XXI.
Fic. 38. Euphalerus nidifex Schwarz. Venationof forewing. See page119.
Fic. 39. Pachypsylla celtidis Riley: Venation of both wings. See page
119.
Fic. 40. Anomoneura mort Schwarz. Venationof forewing. See page 120.
Fic. 41. Pseudococcus citri. Venation. See page 123.
Fic. 42. Dactylopius species. Tracheation of freshly emerged wing. See
page 123. :
Fic. 43. Dactylopius species. Venation. See page 123.
Fic. 44. Aleurodes sp. Tracheation of freshly emerged fore wing. See
page 122.
Fic. 45. Aleurodes sp. Venation of forewing. See page 122.
Fic. 46. Aleurodes sp. Tracheation of hind wing. (Venation is identi-
al with this). See page 122.
ANNALS E. S. A. VOL. 1, Prars sovi.
Sce+R+M+ Cu ee ieee SOO
—— See ioe
Edith M. Patch.
ANNALS, E. S, A. VoL. II, PLATE XVII.
Sce+R+M+ Cu + 1st_A
24 A
13
=
M
Sc: R+M+Cu + 1st_A i
Ediih M. Patch.
ANNALS E. S. A. Vou. II, PLATE XVIII.
Sc+R+M+Cu ,+ (st A
1st A Cu Masa
Edith M. Patch.
ANNALS E. S. A.
VoL. II, PLATE XIX.
R+M Cut 1st A+2d A
Edi.h M. Pateh.
ANNALS E. S. A. VOL. LEP rAnE oxox.
Se-R+M+Cu, + 1st A = aie a tore
Cu,
It,
1st A od A Cuz Mo+4
M+2
Edith M. Patch
ANNALS E. S. A. VOL. II, PLATE XXI.
41 46 .
Edith M. Patch.
ROBBERFLIES OF THE GENUS ASILUS.*
JAMES S. HINE.
The insects of this genus together with a large number of
others belong to the Asilidae which is a family of Diptera com-
monly called robberflies because of their predaceous habits in
the adult stage. These habits usually lead to beneficial results but
none of the species are known to show any indication that they
prefer particular food. They take all kinds of insects that inhabit
like places with them, and have the habit of alighting on a leaf
or a stone, a log or other object, or even on the bare ground to
wait for some unfortunate insect of almost any order to appear,
when they leave their perch and, like a hawk, pounce upon the
chosen victim and at once introduce their piercing mouthparts
and suck away its life blood. Occasionally some observer finds
certain species to be injurious on account of feeding on beneficial
species, and it is no more than likely that all of the members of
the family do some harm. Riley has stated that one of the larger
robberflies may be very destructive to honey bees, and frequently
one may see one or another of the flies with a parasitic insect in
possession. Such habits furnish much of interest to the observer
in the field, and the student who is in search of a subject full of
sentiment will do well in making a selection here.
So far as known the larvae are predaceous and are found in’
dry ground. They work beneath the surface and seek out such
available food as these situations offer. Two larvae of Asilus
sericeus Say, were taken April 3, while spading a garden. They
were located about six inches below the surface of the ground,
color white, naked with the exception of a very few brown hairs
scattered over the body, general form cylindrical, or only slightly
compressed; in appearance much like what in general pass under
the common name of grubs. Total length when full grown 20
millimeters. The pupa stage was reached May 25, and the adult
appeared June 8.
The various species of the family are mostly large insects, but
have been neglected more or less in America, some of the genera
as yet not having received the attention of specialists beyond the
*Contributions from the Department of Zoology and Entomology of the
Ohio State University, No. 32.
136
1909] Robberflies of the Genus Asilus 1
“I
description of such species as have been brought to their atten-
tion for names. There is in the collections of the country, there-
fore, a quantity of undetermined material and not a few undes-
cribed species which are a hindrance to investigators as well as
to the painstaking amateurs whose ideals are to have all speci-
mens named and grouped in reference to some particular system”
under a series of neatly written labels.
The genus here considered is classified in the Asilinae, one of
the four subfamilies into which the family is divided in North
America. In the Asilinae the species are of medium to large
size, long and slender form and _ black, yellowish, brownish or
gray color, with a close general resemblance throughout the
group. The antennae each are composed of three segments of
which the first is longer than the second, and the third longest
of all; the latter segment at its apex bears a long style or arista
composed of two segments of which the first is very short. In
the wing the marginal cell is always closed, as are the fourth pos-
terior and anal cells, and there are either two or three submarginal
cells of different lengths. The members of the genus Asilus are
typical of the subfamily, the antennal arista is naked, the poster-
ior branch of the third vein reaches the margin of the wing be-
yond its apex, there are two submarginal cells in all cases and the
veins which close the discal and first posterior cells are not
parallel.
Ever since Loew monographed the European robberflies in
1848 and divided the genus Asilus of Linneeus into groups with
separate names, students have been at variance as to whether or
not Loew’s names should be considered as genera. In Diptern-
fauna Sudafrica’s issued in 1860, Loew himself raised his names
to generic rank, but Schiner, one of the foremost students of Dip-
tera, writing at about the same date considered Asilus in the wide
sense. Later European students in the main have been inclined
to follow Loew as have some Americans. Lately, however, there
is a tendency to drop Loew’s names and include all of the species
concerned in the genus Asilus, thus following Schiner’s arrange-
ment. Thus Pandelle, an European, writing as late as 1905,
and Williston of this country, in his revised edition of North
American Diptera, published in 1908, have made it plain that
they prefer the latter. Personal inquiry brings out the fact
that nearly all of the prominent dipterologists of the United
States are ready to accept Williston’s views in the matter.
138 Annals Entomological Society of America [Vol. II,
In his Manual Williston says that “‘In general the subdivisions
of the old genus Asilus are very vague and hard to define, and
many of them are doubtfully entitled to recognition. At the
most, few, if any, of them are based upon real generic characters
and the names are only useful as aids in the determination of
the numerous forms.”’
Lundbeck, in his Diptera Danica 1908, II, 53, gives what argu-
ment there is for the other side when he says of these names that
‘Though it cannot be denied that as genera they are far more
closely allied and taken in a narrower sense than genera are com-
monly, I shall yet retain them, chiefly because they are
generally in use and at all events give valuable hints about the
natural classification of the species.”’
Before beginning the study of the species considered in this
paper, I procured representative specimens of Loew’s groups
from Europe by exchanging with Dr. M. Bezzi and others, and
with these for reference have undertaken to place the North
American species. The list offered below indicates the success
achieved, but as may be noted some of the species do not fall
naturally into any of the groups and even in some cases where a
decision appears to have been reached, there are difficulties in the
way of a perfectly satisfactory conclusion. It may be mentioned
also that Dr. Bezzi encountered some difficulty in placing the
Nearctic species, for at the end of the Asilinae in the recent Cata-
logue of Palearctic Diptera is a lst of no less than twenty-nine
species, which from the evidence cannot be placed in the so-called
genera, but are given as falling under the genus Asilus in the wide
sense.
By giving only Asilus generic rank and using the other names,
that often have been considered as genera, simply as names of
groups, some questions in priority come up for solution. Since
Europeans were the first to arrange their species in reference to
Loew’s classification, the proper procedure would be for them to
be the first to discard it. As it is students in America have
described many of their species on the ground that Loew’s names
were valid generic terms, using the same specific name in two or
more genera. If these names are considered as genera in Europe,
as is the case in the recent catalogue mentioned above, and only
as groups of one genus in America, a student in the latter country
hardly knows what is the proper disposition of the synonymy
question in his fauna. With these facts confronting me, I do
1909] Robberflies of the Genus Asilus 139
not know of a better way of accepting Williston’s views on the
extent of the genus Asilus than to restrict it only by the descrip-
tion I have given above, and this would also agree with Schiner’s
views so far as European species are concerned. Having reached
this conclusion I have worked out the synonymy accordingly,
so far as evidence could be obtained, and it develops that several
of the names that have been used as names of species in America
become untenable on account of having been used earlier as names
of species in the Old World.
Macquart’s studies in the subfamily in general preceded those
of Walker, but since the former writer used a number of names
that have been employed by older writers, he made some syno-
nyms which must give way to names which Walker gave to the
same species, and a similar condition exists in reference to the
work of the later authors.
In his treatment of the Asilinae Lundbeck has followed Mik
in reference to the method used in referring to the color and the
arrangement of the bristles of the legs. | When these organs are
stretched out at right angles to the body, the side which looks
forward is anterior. As this answers all purposes and is in use
it is adopted in this paper.
I have received encouragement in this work from most of
the dipterists of the United States and take this opportunity
to express my appreciation of their good will. For the loan of
specimens, without which I could not have hoped to attempt this
paper, I am under obligations to the following museums and
private parties: The United States National Museum, including
an abundance of material collected by Coquillett on the Pacific
Coast; the Francis Huntington Snow Collection of Kansas Uni-
versity; the collection of the Massachusetts Agricultural College
and the collection of the University of Dllinois. Chas. Dury,
Chas. W. Johnson, Dr, O. A, Joaannsen, Dr. C. F. Adams, Dr.
W. A. Nason, Franklin Sherman, Jr., Dr. O. S. Westcott, 5S. A.
Rohwer, H. S. Harbeck, and Dr. Nathan Banks have loaned
or given me specimens from their own private collections
or from the collections of the institutions which they represent.
Furthermore Dr. W. D. Hunter, of the Bureau of Entomology,
Washington, D. C., has sent for study a large collection of Asilinae
collected during the time the studies of the cotton-boll weevil
and other species have been in progress in Texas.
140 Annals Entomological Society of America [Vol. II,
LIST OF THE SPECIES.
Group Asilus.
sericeus Say, Jour. Acad. Sci. Phil., III, 48. Compl. Works II, 68. Wiede-
mann, Auss. Zw. I, 429. Williston Trans. Am. Ent. Soc. XI, plate II,
figure 10. Kan. Univ. Quar. II,68. Riley Mo. Rept. JI,123. Harris
Ins. Inj. Veg. 605.
herminius Walker, List II, 410.
midas Brauer, Sitzungsb. Kaiserl. Akad. Wissensch. XCI, 387, plate II, figure
1. Williston, Kan. Univ. Quart. II, 69. Osten Sacken, Biologia,
Diptera I, 209.
Group Rhadiurgus.
leucopogon Williston, Kans. Univ. Quart., II, 75.
cacopilogus new species.
Group Philonicus.
arizonensis Williston, Kan. Univ. Quart. II, 76.
fuscatus new name.
obscurus Hine, Ohio Naturalist VII, 117. Name preoccupied.
limpidipennis new species.
arizonensis as determined by Hine, Ohio Naturalist VII, 116.
rufipennis Hine, Ohio Naturalist VII, 117.
truquii Bellardi, Saggio II, 52. Van der Wulp, Tijdischr. v. Ent. XXV, 116.
* Williston, Biologia, Diptera I, 328.
taeniatus Bellardi, Saggio II, 55. Synonymy by Williston.
plebeius Osten Sacken, Biologia, Diptera I, 209. Synonymy by Williston.
tuxpanganus Bellardi, Saggio appendix 22. Hine, Ohio Naturalist VII, 118.
Group Nigrasilus.
nitidifacies Hine, Canadian Ent. XL, 202.
Group Antiphrisson.
astutus Williston, Kans. Univ. Quart. I], 70.
Group Eutolmus.
lecythus Walker, List II, 451.
femoralis Macquart, Diptera Exot. Supplement II, 45.
Group Machimus.
avidus Van der Wulp, Tijdschr. v. Ent. XII, 82.
occidentalis new species.
tenebrosus Williston, Biologia, Diptera I, 328.
griseus Hine, Ohio Naturalist VII, 29.
Group Neoitamus.
affinis Williston, Kan. Univ Quart. II, 73.
coquillettii new species.
brevicomus new species.
orphne Walker, List II, 456.
auceps Van der Wulp, Tijdschr. v. Ent. XII, 84.
aeneobarbus Loew. This name was sent to Osten Sacken in a letter and
so far as I can find, no description was ever written.
distinctus Williston, Kansas Univ. Quart. II, 73.
terminalis new species.
flavofemoratus new name.
flavipes Williston, Kansas Univ. Quart. II, 72. Name preoccupied.
Group Stilpnogaster.
auriannulatus Hine, Ohio Naturalist VII, 29.
Group Heligmoneura.
angustipennis new species.
latipennis new species.
1909] Robberflies of the Genus Asilus 141
Group Cerdistus.
willistoni new name.
angustifrons Williston, Kansas Univ. Quart. II, 71. Name preoccupied.
auricomus new species.
piceus new species.
montanus new species.
albicomus new species.
lepidus new species.
Group Tolmerus.
antimachus Walker, List II, 454.
callidus Williston, Kansas Univ. Quart. II, 75.
delusus Tucker, Kansas Univ. Sci. Bull. IV, 92.
johnsoni new species.
maneei new species.
notatus Wiedemann, Auss. Zw. I, 451. Williston, Kansas Univ. Quart. II, 74.
alethes Walker, List II, 454.
nove-scotiae Macquart, Dipt. Exot. Suppl. II, 46. Williston, Kansas Univ.
Quart. II, 68.
paropus Walker, List II, 455.
prospectus Tucker, Kansas Univ. Sci. Bull. IV, 94.
prairiensis Tucker, Kansas Univ. Science Bull. LVE29 3).
annulipes Macquart, Dipt. Exot. I, 2, 149. Name preoccupied.
sadytes Walker, List II, 453.
tibialis Macquart, Hist. Nat. Dipt. I, 313. Name preoccupied.
snowii new name.
annulatus Williston, Kansas Univ. Quart. II, 70. Name preoccupied.
Group Epitriptus.
erythocnemius new species.
Group not determined.
gracilis Wiedemann, Auss. Zw. I, 445. Osten Sacken, Catal. 235, note on the
type.
auratus Johnson, Proc. Acad. Nat. Sci. 1895, 305.
californicus new species.
mesae Tucker, Kansas Univ Sci. Bull. IV, 92.
rubicundus new species.
KEY TO THE SPECIES
1. On each side of the abdomen near the incisures with a transverse
row of bristles which are clearly larger than the hairs to be seen
clearly larger than the hairs of the other parts of the segments.... 2.
Large, bright colored species, wings distinctly colored all over...... 3.
Smaller, modest colored species, wings pale colored or hyaline...... 4.
3. Body and legs uniformly black, wings uniformly rich yellowish
DOWIE, Ricoh cece soe encsleeege een see ohare oe weaned Seite aire ance ee. midas Br.
Body brown, legs pale brown, wings brown with the veins margined
With paler! faeiamrstecne vers wee eiensea eter ck seeer susie acres ¢ sericeus Say.
4. Scutellum with more than a dozen long bristles on its margin, female
genitalia with a circlet of strong spines at the tip.............. 5.
Scutellum with not more than half a dozen long bristles, female geni-
talia not wiuoancincletohispinesraitebWentip rier cas ce crs se 1 6.
5. Upper forceps of the male genitalia protruding half their length
beyond! the lower’ forcepsies yen ees 8 wns alee sees leucopogon Will.
Upper forceps of the male genitalia about equal in length to the
KOR org nes omen aaatinis Soot GO claties sicMcnanae deo coins as hace cacopilogus n. sp.
6. Body black, space between the antennae and the facial gibbosity largely
shininotblack pacity noe ic ee Ee on he Sos nitidifacies Hine
Body brown, face entirely clothed with yellowish dust...... astutus Will.
to
142
10.
Tike
13.
14.
16.
18.
19.
Annals Entomological Society of America [Volvii,
. Ovipositor short, conical, with four spines on the upper side at the
apex, upper forceps of the male genitalia excavated on the apical
part so that when viewed from above there is a distinct open space
between them... ..09.5 0 s0g.6+4.6 02.0 «230 © See ne nenenenen sie eek 5
No spines at the tip of the ovipositor.) >). eee een) 13.
. Legsin large part reddishi no. wv... + 2. 4 ee eee ets cree 9.
At least the femora of all the legs black. -- amet cee ce 12:
. Largest species falling under 7, wings smoky, legs dark red, abdomen
pure black’ banded, with gray...:.... see eee arizonensis Will.
Smaller species, wings hyaline or reddish, abdomen gray black band-
ed: with gray: cc5 8 oe eas ce ns oe 0 oe Oe et tey a ne ls 10.
Wings limpid, legs pale reddish, brown of fuscus in places.........
limpidipennis n. sp.
Wings reddish, legs pale reddish, usually black in places.......... hile
Legs rather slender, anterior side of each femur in large part black,
upper forceps of the male genitalia with the angle at the beginning
ot the excavation produced inward and backward so that it meets
and crosses its fellow of the opposite side................. truquii Bell
Legs rather thick, each femur black except at the tip of the anterior
side, angle of ‘the upper forceps not produced........ rufipennis Hine
. Legs, except the extreme bases of the tibiae, black..... tuxpanganus Bell.
Tibiae; and tarsi inslarge part reddisitiaccs. es See fuscatus n. name.
Arista of the antenna about twice as long as the third antennal seg-
ment, legs almost emtirely: pale reddish...2-. 0. -1ee- oe gracilis Wied.
Arista at most only a little longer than the third antennal segment,
legs Vusually jparthy “black-tie cei a eeu eye renee mel eee 14.
Four or more bristles on the margin of the scutellum, ground color of
the abdomen, except im one species, clear black: 2.7 .-........ iG
Normally two bristles on the margin of the scutellum. These bristles
may be variable to the extent of having one or two additional
smaller ones in an occasional specimen of a species.............23.
. Wings glassy hayline, legs in part bright yellow, Sixth and seventh
segments “of the female abdomen shining black, that is, included in
the ONDPOSTLOTE 1A ohewen sect Pease ke tenes ocelot ane atis se Gra ian sl Guarepeats 18.
Wings partly brownish, legs in part pale reddish. Sixth and seventh
segments of the female abdomen not included in the ov ipositor. .16.
Dark gray species with gray posterior margins to the abdominal seg-
ments. Posterior margin of the eighth ventral abdominal segment
produced and furnished with a dense tuft of coarse hairs......
occidentalis n. sp.
Black species. Posterior margin of the eighth ventral abdominal
segment not produced and not furnished with a tuft of hairs....17.
. Shining black species with golden posterior margins to the Aatlossete
nal ‘segments Bat cnt revs ertagahs 2 Oc As eine hap og sede OnaE auriannulatus Hine
Rather dull black species with gray posterior margins to the abdomi-
PCH SE OTM EMIS iano aieede: oo ose ecu Naan aagets ovo eere ee a ee callidus Will.
Hairs of the face, mystax, bright yellow. Male forceps notched
on ithesupper side at the apexes 1. ce oc eee ee aoe 19
Mystax wholly or in part black or white, or else the male FORCEDS
not notched on the upper side atthe apex. jee ee ee 20.
Altsthe femroravblack® <5... tse a se i eee eee orphne Walker
Front and middle femora almost entirely yellow............. affinis Will.
. Front and middle femora yellow, each with a black stripe on the
WP PSRSIG eA esgic ysuevere sca hedoce Dace ara SEE ee eae flavofemoratus n. name
All the femora black, at most with only a preapical band yellow... .21.
. Male forceps notched on the upper side\at the apeéxe. ©... 30. .5.- 4... 22.
Male forceps not notched on the upper side at the apex. ...terminalis n. sp.
1909] Robberflies of the Genus Asilus 143
22. Dorsum of the thorax for its entire length with a crest of long
aE ee errs OEE Te EG Act ene, cond eas ais coquillettii n. sp.
Dorsum of the thorax with the hairs on the anterior part distinctly
shorter thant omstRempOsteriOG Parts genic. 6 ace. 6 occu oc brevicomus n. sp.
23. From dorsal view male forceps wider at two thirds their length than
at their base. First segment of the female ovipositor, that is, the
eighth abdominal segment longer than the sixth and seventh
TUMITE GE Mien eeee HRC Mey eho enna ccna ca-° paleo len ed) een et esi dees ye 24.
From dorsal view male forceps gradually narrowed from near the
base to the apex. First segment of the female ovipositor of nor-
mal length, not so long as the sixth and seventh segments united. .31.
24. Male forceps distinctly wider than the abdomen at the middle of its
ene ae ee ae
Male forceps not wider than the abdomen at the middle of its length. . 26
25. Wings narrow, male genitalia more than two millimeters in width
ate Che * a De xsi ieee crate cee ee oe ce ree seus angustipennis n. sp.
Wings wide, male genitalia less than two millimeters in width at
NE SUC trap tse Per rt eee eae RMSE, oT Get ah atone ok latipennis n. sp.
26. Mystax golden, usually with a few black bristles above. Preapical
bristles of the abdominal segments pale yellow, male wings not
distinetlyeiprown, atsmmddile 2yo6 2). 6 10s s4 terme ese. auricomus n. sp.
Mystax gray with some bristles above, or if the mystax is not gray,
the wings of the male are distinctly brown at middle, preapical
bristles of the abdominal@segments) eray e242. ..500:8 0c os eile
. Wing of the male distinctly brown at the middle of the length...... 28.
Wing in both sexes hyaline, or very faintly tinged with reddish all over29.
28. General color black. Viewed from above the male genitalia on each
sidemeventya cunvyedetomunemapexamrs ane ae 4 ier netsh: lepidus n. sp.
General color brown. Viewed from above the male genitalia on
each side quite suddenly contracted just before the apex.......
willistoni n. name.
29. Facial gibbosity small, not reaching half way to the antennae, wings
Bisel HG Siaeynysg cick egies, « a atheattel soph Soy eg Bush home oketien tae | montanus n. sp.
Facial gibbosity large, reaching more than half way to the antennae. .30.
30. Body gray, legs red, wings faintly tinged with reddish, bristles and
iw)
|
airs On thesbodyaawihiten. asain Unaaese cond. seine an 36 albicomus n. sp.
Body black, legs dark, wings very faintly smoky, bristles of the
Wetopreb.< have Tees canosThy WICK. oc oclc co Goons noo odso bb don piceus n. sp.
31. Wings clear hyaline, rather large species, upper forceps of the male
genitalia splthrat Mune tip ce cee tenet balls 5 le desde oi californicus n. sp.
Wings washed or spotted with gray at the apex and along the poster-
ior border, species of varia ble sizes, upper forceps ‘of the male
genitalia entirerat bier Pex amiss wu sian reed ss See AA Sas 32.
32. Very small species; antennal arista very short, not over one fourth
the length of the third segment and not distinctly differentiated
rosea Habs} eAdae Mite wo mdlom Wo dia bioo.c Gola 6 pboleteo oa Hole o CASE mesae Tucker.
Larger species, antennal arista distinctly differentiated from its
Selena alta man cecuotho ain 4.0 Ove id Slate a Rea nONs Ole D Bia ole: cle eae ens Eee RENO. tT
33. Arista of the antenna about as long as the third segment, only two
bristles in addition to the apical. bristles on the front side Of the
hind tibia. These bristles are not entirely uniform but may be
aKeyeresareleyel qoyororal SWAMI. - sec con okoapoecuonogoouu Munn PEesone oot D4.
Arista of the antenna usually not over two thirds as long as the
third segment, three bristles on the front side of the hind tibia... .44.
144 Annals Entomological Society of America [Melani
34. Gray of the wings in the form of spots in the cells at the apex and
along the posterior border, leaving the veins margined with
Inyalane: "cso aie ah seaiceph cucsavsreveunteee ate cheer oy no ees aang nee tenebrosus Will.
Apex and posterior border of the wing uniformly gray. In some
Species this colormne: is) very fein... sen eee ee 30.
35. Ventrally the posterior margin of the eighth abdominal segment of
the male is: somewhat produced’... j...4 552 ame avidus v. d. Wulp.
Ventrally the posterior margin of the eighth abdominal segment is
NOt PFOGMCEG oc. Fe He rouncce Cn sts a ee eee ae a 36.
36; Black species fiat ty geiemsnae ete eda oe ee c enc ee ent Reon eet Ye oe SH
Gray; brown, or yellowish brown species: 6 )...).stn- dace oe 40.
37. Small species, legs black, at most with just a trace of reddish on the
baselofiveach' tibiae ye. sence ee eae Oe eae maneei n. sp.
Largerispecies, WQegs plainly marked with red... 145:.+22,05 400 38.
38. Male upper forceps long, usually longer than the last three abdominal
segments together. Mystax largely black........... sadytes Walker
Male upper forceps shorter, usually about as long as the first two
abdominal segments together. Mystax largely light.......... 39.
39. Eastern species. Each tibia red on basal third............. notatus Wied.
Western species. Each tibia largely red, especially on the posterior
SIS 10 a chert fed, «nee He cae bee Cent a Oe een nee See callidus Will.
40. Rather large species, wings wide, end lamellae of the ovipositor
wedged)! hss. o1.sa7a Jame one | Melee eur aes Coches cee ae lecythus Walker
End lamellae of the ovipositor not wedged in...................:- 41.
Ale Alle the femora, whollyeiblackwatasn sess ae ae novae-scotiae Macquart
Femora partly red, at least each with a preapical red band........ 42.
42. Femora variable, sometimes wholly red, commonly with only the
posterior side red, mystax almost always white or yellowish white. 43.
Femora largely black, mystax in large part black, from side view
male genitalia widest at two thirds the length...... erythocnemius n. sp.
43. Small reddish species, mystax straw yellow, legs mostly red, male
Senitalia rede Moen toys ee enn Cn anche cee hare eee rubidus n. sp.
Larger species, mystax white, occasionally a few black bristles above,
male.centtalia plackey esa eran arene ene ene antimachus Walker.
44, Small reddish species, legs red, at most only the anterior side of each
femur slightly darkened, male genitalia red........... rubicundus n. sp.
Femora in large part black. Species of ordinary size.............. 45.
45. Each femur with a preapical band and the posterior side red....... 46.
Bach femur with onlyratpreapical bandi rede. =e ae nee Orie 48.
46. Dull gray species, under side of each front femur with numerous
pray IH atS; © Nie wats eee tock eres Aaa CRC Eee delusus Tucker
Bright gray or yellowish species, under side of each front femur with
a longitudinal TOW Of TIStlest 7ir.33: Geile eee wateannre te Cera eee eee 47.
47. Thorax with a very distinct mid-dorsal black stripe which reaches its
entire length, male senitala darksrede i 4-0 5° 5 ee johnsoni n. sp.
Mid-dorsal thoracic stripe not very distinct and abbreviated behind.
Male genitalia black, or at least dark fuscus....... prairiensis Tucker.
48. Under side of each front femur furnished with abundance of rather
lonig hairs, not TOGO} d= aR ek AM ee a Allan top Ars snowli n. name.
AQ, eis on the under side of each front femur white and rather weak.
Upper forceps of male genitalia from side view widest at two thirds
themlensthigeact- tat ee eRe een eo ae erythocnemius n. sp.
Bristles on the under side of each front femur rather stout, nearly
always partly or wholly black, upper forceps of the male genitalia
from side view of the same width thr OUGHO UGH as eee paropus Walker
1909] Robberflies of the Genus Asilus ards
Asilus gracilis Wiedemann. (Figs. 1 and 2.)
A long slender species with yellow body and legs, facial gibbosity
very small, mystax composed of a few white bristles. Length 15-17
millimeters.
Head rather large, face and front narrower than usual; palpi and
proboscis dark, the former clothed with light hair; first two segments
of the antenna yellow, third brown, first segment longer than the
second, arista near twice as long as its segment; occipito-orbital bristles
largely yellow, although in some specimens these are nearly all black.
Facial gibbosity hardly evident, mystax composed of a dozen or so
light colored hairs and bristles. Thorax yellow, thinly clothed with
gray dust, mid-dorsal stripe dark brown; wings hyaline with a slight
clouding at the apex and along the posterior border; legs pale, anterior
side of each femur, apex of each tibia and of each tarsal segment
usually darker, although there is quite an amount of variation in the
color of the legs. Abdomen somewhat darker than the rest of the
body, nearly uniformly colored but with narrow band on the posterior
border of each segment a little lighter; these narrow bands are each
preceded by about four yellow bristles on either side of the abdomen.
Male upper forceps short, rounded at the tip, of nearly the same
width throughout; lower forceps dropped so that there is always a
distinct space between the upper and lower; end lamella of the oviduct
style-like.
Specimens from North Carolina, Texas, Florida, and Georgia.
The type of Asilus auratus has been studied and is this species with-
out question. The long antennal arista makes the species easy of
identification.
Asilus lecythus Walker (Figs. 3 and 4).
Asilus femoralis Macquart is this same species but as there is an
older Asilus femoralis by Zeller from Europe, Isis 1840, 49, 3, Walker’s
name must be used.
Color yellowish brown; femora largely black but each with pre-
apical band and often with the posterior side yellow, tibiae and tarsi
in large part yellow. Length 18-21 millimeters.
Face and front covered with pale yellowish dust; facial gibbosity
prominent, mystax composed of a few black hairs above and numerous
pale yellowish hairs below, beard pale, palpi and proboscis black,
the former with light colored hairs; antennae largely black but apex
of second and base of third joints plainly yellow: first two segments
together equal in length to the third which is of nearly the same length
as its arista. Occipito-orbital bristles black and yellowish, somewhat
variable; other bristles and hairs of the occiput light colored. Thorax
dull gray on the sides, more yellowish above, mid-dorsal brown stripe
divided anteriorly; wings largely hyaline, apical portion and posterior
border slightly clouded; coxae colored like the sides of the thorax,
femora variable, in some specimens black with the exception of the
preapical band; in others, the whole posterior side yellow, even the
different femora of the same specimen may vary in coloration; tibiae
146 Annals Entomological Society of America [Viol iT.
yellow each with a narrow black apex, metatarsi yellow apex of each
black; other tarsal segments black mostly with yellow bases. Abdo-
men dark brown in ground color, thinly clothed with yellowish dust;
posterior margin of each segment plainly lighter colored than the
other parts and preceded on each side by a few bristles. End lamella
of the oviduct wedged in.
This is the only American species known to me with the end lamella
of the oviduct wedged in. Specimens from Pennsylvania, New Jer-
sey, Maryland, North Carolina, Massachusetts, Maine, Ohio.
Asilus tenebrosus Williston (Fig. 5).
Machimus griseus Hine, The Ohio Naturalist VII, 29, isa synonym.
There is an older Asilus griseus Wiedemann from Java, Dipt. Exot.
11925 i:
Male and female brownish gray with reddish legs and narrow white
margins at the apexes of the abdominal segments. Wings hyaline with
well defined dark markings in the cells at the apex and along the pos-
terior margin. Length 17 to 20 millimeters.
Front and face of usual width, facial gibbosity prominent extend-
ing two thirds of the distance from the oral margin to the antennae;
mystax black above and white below; antennae black, rather long,
third segment of each with its style decidedly longer than the other
two segments combined, style about equal in length to the remainder
of the segment, beard white. Thorax dark in ground color but
clothed with gray dust which is denser in some places than in others,
thus giving the part a variegated appearance; a wide mid-dorsal black
stripe divided anteriorly by a narrow grayish interval; scutellum with
two black bristles at the apex; legs red with numerous black bristles,
coxae black, each femur dark on the anterior side except just before
the apex; wing clear hyaline with dark markings as follows: marginal,
both submarginal and first posterior cells each with a stripe which
reaches the apex of its respective cell; second posterior, discal and
anal cells each with an angular spot not contiguous with the margin
of the wing; halteres pale yellowish. Abdomen of the same general
color as the thorax, with a light colored annulation preceded by a row
of white bristles at the apex of each segment. Eigth segment of the
male distinctly produced below, but not with an appendage as in
some of the European species of the group; male genitalia reddish in
color, oviduct shining black.
Several specimens from southwestern Colorado and Huachuca
Mountains, Arizona; Flagstaff and Williams, Arizona, collected by
H. S. Barber; White Mountains, New Mexico, by Townshend; Pecos
and Beula, New Mexico, the latter place at an elevation of 8000 feet,
by Cockerell.
The hyaline wings with the dark colored markings in the cells
as described and the wide male genitalia are sufficient to separate
the species from its near relatives.
T909| Robberflies of the Genus Asilus 147
Asilus avidus Van der Wulp (Fig. 6).
General color reddish gray, abdomen darkest and with prominent
gray margins to each of the segments above; eighth ventral segment
of the male slightly produced posteriorly but with only a few bristly
hairs on its margin. Preapical band and usually the posterior side
of each femur and the tibiae and tarsi dark reddish. Length 15-18
millimeters.
Facial gibbosity rather prominent. With black bristles above and
yellowish gray bristles below; each antenna black, third segment exclu-
sive of the arista scarcely as long as the other two, arista hardly as
long as the third segment: beard white; occipito- -orbital bristles yel-
lowish gray with the exception of three to five black bristles behind the
upper part of each eye. Thorax gray with the usual dark markings
above, bristles of the posterior part prominent and black, about four
black bristles on the margin of the scutellum. Legs clothed with
fine hair and prominent bristles; the hair is either white or yellowish
but the bristles are black; each coxa and the anterior side of its femur
black, otherwise each leg is reddish, slightly darker in some places
than in others. Wing hyaline with a slightly darkened tinge at the
apex. Abdomen rather dark but appearing yellowish gray from cer-
tain angles; narrow posterior margin of each segment plainly gray,
margin of the eighth segment below in the male plainly produced and
forming a distinct angle but not with so prominent a tuft of hairs as
in occidentalis.
Several specimens from Copeland Park and North Boulder Creek,
near Boulder, Colorado, the latter part of August and the first part of
September, 1907. Collected by S. A. Rohwer and Glen M. Hite. Also
from Beula, New Mexico, 8000 feet elevation, by Cockerell.
Everything considered this is most like Van der Wulp’s species.
The tibiae are most plainly red at base but nowhere is there a sharp
differentiation of colors as in most specimens of occidentalis. The
male genitalia are nearly straight, and in this respect quite different
from those of occidentalis. The latter character more than any
other has led to the determination of this species as avidus.
Asilus occidentalis n. sp. (Fig. 7).
General color gray with the thorax often yellowish gray. In some
specimens each femur has a preapical reddish band, in others this is
not the case. Posterior margin of the venter of the male eighth ab-
dominal segment furnished with a dense cluster of rather long bristly
hairs. Length 14-18 millimeters.
Facial gibbosity prominent, mystax largely black but with a num-
ber of yellowish hairs below; face clothed with yellowish dust, beard
white; each antenna black with the third segment exclusive of the
arista slightly longer than the first two segments together, arista
scarcely as long as the third segment. Occipito- orbital bristles mostly
black, but in some specimens ‘nearly all yellow. Thorax gray or yel-
lowish gray, median darker markings not especially prominent, scutel-
lum with at least four bristles on its margin. Legs dark, each femur
148 Annals Entomological Society of America [Vok. EL;
may or may not have a preapical reddish band; each tibia plainly
reddish at the base and this color is not always well defined below
and may extend nearly the whole length, especially on the upper side;
tarsi black although the metatarsi are often largely reddish; wings
hyaline with the apex and the inner margin of. each faintly gray.
Abdomen dark, differing slightly according to the angle from which
it is viewed, posterior margin of each segment plainly gray.
Specimens are at hand from British Columbia collected by Harvey,
Venables and R. S. Sherman; Ormsby County, Nevada, by C. F. Baker;
several counties in southern California by Coquillett; also from Eldo-
rado County, California, and from Oregon and Washington.
The fact that the male genitalia are bent upward near the middle
as shown in the figure, serves to designate the species. It is much like
Asilus notatus of the eastern states but in the male the extension of
the ventral margin of the eighth abdominal segment with its dense
tuft of long hairs, is distinctive and in both sexes the form of the third
antennal segment is a useful character. In occidentalis this antennal
segment is plainly wider than in notatus.
Asilus auricomus n. sp. (Figs. 8 and 9).
General color yellowish gray, mystax largely composed of golden
yellow bristles, also, yellow bristles before the incisures of the abdomi-
nal segments. Legs yellow with an elongate black marking on the
anterior side of each femur and a black spot at the apex of each tibia.
Length 15 millimeters.
Front and face rather narrow and covered with yellowish gray dust.
Facial gibbosity rather prominent, reaching half way to the antennae,
furnished with yellow bristles intermixed above with a few black ones.
Beard gray, occipito-orbital bristles yellowish gray. Antennae black,
third segment of each exclusive of the arista about equal in length
to the other two; arista decidedly longer than the third segment.
Thorax yellowish gray with the usual dark markings; scutellum
clothed all over with rather long yellow hairs and with two yellow
bristles on the margin. Wings hy aline on the basal part, and grayish
at the apex and along the inner margin. Legs largely shining yellow;
coxae dark, clothed with gray dust and yellow hairs; each femur
with an elongate black marking on the middle of the anterior side;
each tibia with a black spot at the apex of the anterior side; tarsi
infuscated, especially on the apical part of each segment; pulvilli
yellow. Abdomen yellowish gray with yellow hairs on the segments
and several yellow bristles on each side before the incisures. Male
appendages, viewed from above, widest near the apex; each appendage
black on the basal part and yellowish distally.
Several specimens of both sexes from Medina County, Ohio, taken
in August. Others from Iona, N. J., collected by E. Daecke, New
Haven, Connecticut, by R. L. Butrick, southern Illinois by C. A.
Hart, and from Germantown, Pa.
1909] Robberflies of the Genus Asilus 149
Asilus piceus n. sp. (Figs. 10 and 11).
Nearly black species with hyaline wings and brown legs. Length
15 to 17 millimeters.
Facial gibbosity not very prominent, reaching half way to the
antennae, mystax composed of black and white hairs, intermixed,
beard white, occipito-orbital bristles largely black, face and front
clothed with gray dust, antennae black, third segment exclusive of the
arista near the length of the other two, arista equal to the third seg-
ment in length. Thorax brown in ground color, clothed with gray
or yellowish gray dust, mid-dorsal stripe rather wide, black, split on
the anterior part and abbreviated behind; wings clear hyaline in both
sexes, veins brown; legs brown with light colored hairs and with light
and dark bristles. Coxae colored like the thorax, anterior sides of
all the femora, apexes of all the tibiae and the last four segments of
each tarsus darkened more or less. Abdomen black, posterior margins
of the segments narrowly gray, male forceps distinctly excavated at
the upper corner of the apex, first segment of the oviduct nearly as
long as abdominal segments five, six and seven; apical segment style-
like and near the length of abdominal segment seven.
Specimens from Amherst, Massachusetts, from the collection of
the Massachusetts Agricultural College.
Asilus montanus n. sp. (Figs. 12 and 13).
Rather small, slender, gray species with hyaline wings, largely
reddish legs, very small facial gibbosity and mystax composed of a
few bristly hairs which are mostly white, although there are a few
black ones above. Length 13 millimeters.
Face clothed with gray dust; palpi and proboscis black, the former
with white hairs; occipito-orbital row of bristles with a few black
ones back of each eye, otherwise all the bristles and hairs of the occi-
put white. Thorax gray, mid-dorsal stripe black, divided before and
abbreviated behind; several black bristles on the posterior part, two
black bristles on the margin of the scutellum. Wings clear hyaline;
legs reddish, anterior side of each femur, apex of each tibia and each
tarsal segment fuscous. Abdomen clothed with grayish dust, most
pronounced on the posterior margins of the segments. These margins
are preceded on each side by two or three small bristles. Forceps of
the male shining brownish black, each upper with a distinct angle on
the dorsal side at about four fifths of the length.
This species is entirely distinct from its relatives on account of its
_small facial gibbosity and the few bristles composing the mystax.
The male genitalia are distinctive also.
A male collected by Cockerell at Alpine Tavern, Mount Lowe,
California, at an altitude of 5,000 feet. Taken August 12th. Type
in the United States National Museum.
150 Annals Entomological Society of America [Vole tr
Asilus albicomus n. sp. (Fig. 14).
Medium sized gray species with red legs and hyaline wings in both
sexes. Bristles of the whole body, except a few on the feet and in the
extreme upper part of the mystax, white. Length 13 to 15 milli-
meters.
Facial gibbosity prominent reaching nearly half way to the anten-
nae. Mystax composed of many light hairs, only a few dark ones
above; bristles and hairs of the occiput light colored; thorax gray with
the usual dark markings above; wings hyaline in both sexes; legs red,
femora a little darker anteriorly than posteriorly; each abdominal
segment with a gray posterior margin preceded by a row of white
bristles which reach nearly over the dorsum, especially forward.
Both sexes taken by H. K. Morrison in Montana. Type male and
female in the United States National Museum.
Asilus lepidus n. sp. (Figs. 15 and 16).
Dark colored species, facial gibbosity prominent, mystax composed
of numerous hairs, femora largely black, especially on the anterior
sides; tibiae mostly reddish, wings in the female hyaline, in the male
infuscated, especially on the posterior half. Length 15 millimeters.
Facial gibbosity prominent, mystax composed of numerous hairs,
black above, light below; antennae black; thorax clothed with gray
dust leaving the usual markings on the dorsum nearly bare, black bris-
tles on the posterior part above, scutellum with two black bristles on
its margin. Wing in the female clear hyaline, in the male somewhat
infuscated especially beyond the branching of the second and third
veins, and somewhat along the veins of the basal part of the wing.
Legs with femora black anteriorly and dark reddish posteriorly, tibiae
dark reddish, apex of each darkest, metatarsi largely red, other tarsal
segments mostly fuscous. Abdomen black, each segment above with
a prominent gray border which is preceded by a row of bristles which
nearly reaches over the dorsum of the insect, especially forward.
Male from Colorado. Female from the White Mountains of New
Mexico, about 6800 feet elevation, collected July 23, by Townsend.
Types in the United States National Museum.
Asilus willistoni n. n. (Figs. 17, 18 and 19).
New name for Asilus augustifrons Williston, which is preoccupied
by Asilus augustifrons Loew from Asia Minor, Linn. Ent. (1849) IV,
126, 64.
Rather large grayish brown species with brown legs and narrow
front and face, wings of the male more or less dark from the base of the
first submarginal cell, of the female nearly clear hyaline. Length
15 to 20 millimeters.
Head of medium size; front and face narrow, clothed with dust
which varies in different specimens from gray to golden, facial gib-
bosity prominent, mystax composed of a large number of hairs, which
are black above and white, or sometimes golden, below. Antennae
black, first two segments nearly equal in length and together about as
long as the third which is of nearly the same length as the arista.
1909] Robberflies of the Genus Asilus I51
Beard gray; occipito-orbital bristles gray as are all the hairs of the
occiput. Palpi and proboscis black, the former with gray hairs.
Thorax gray, often with a yellowish tinge above, mid-dorsal stripe
black. Wings yellowish in the male quite dark from the base of the
first submarginal cell, especially along the costal border; in the female
the wings are much lighter than in the male and in some specimens
these are practically hyaline. Legs brown, femora somewhat variable
but usually distinctly darker anteriorly, tibiae darkened at the apexes,
tarsal segments mostly dark, but the first on each leg lighter except
at the apex. Abdomen brownish black; on the posterior margin of
each segment is a white band which is preceded by a row of white
bristles which are largest anteriorly. Male forceps black and shining,
upper forceps swollen, thickest beyond the middle of their length,
oviduct long and slender but not including the seventh abdominal
segment.
The description and drawings were taken from the type specimens
which are in the Francis Huntington Snow collection of the University
of Kansas. Other specimens from western Washington collected by
H. K. Morrison,:-Vernon, B. C., by E. P. Venables, Goldstream, B. C.,
by R. V. Harvey.
Asilus auriannulatus Hine.
General color shining blue-black with uniform pale brown wings and
black and yellow legs. The second, third and fourth segments of the
abdomen each with a golden yellow annulus at the apex. Length
14 to 17 millimeters.
Front and face rather narrow, the latter covered with golden
yellow pollen between the callosity and the antennae, callosity slightly
elevated, mystax black, beard white, third segment of the antenna
about as long as the other two together, style shorter than the remain-
der of the segment; thorax dark in ground color, clothed with pollen
which is denser in some places than in others, mid-dorsal stripe opaque
black, narrowly divided on the anterior part; scutellum with several
black bristles at its apex; wing uniformly pale brown all over with a
shght intensity of coloration on the margin of the second vein near the
middle of its length. Legs black and yellow, a preapical ring on each
femur, all the tibiae except at apexes and bases of the tarsal segments
yellow, other parts black; hind femora somewhat variable in that the
yellow is likely to increase at the expense of the black; halteres yellow.
Abdomen shining blue-black, second, third and fourth segments each
with a golden yellow annulus at apex not preceded by a row of bristles;
eighth segment below not widened but furnished with a conspicuous
tuft of erect hair. Genitalia of both sexes shining black, of the male
somewhat wider than the abdomen when viewed from above and
about as long as the seventh and eighth segments combined.
Several specimens of both sexes taken in the Hope Mountains of
British Columbia by R. V. Harvey and R. S. Sherman of Vancouver,
during ‘the first part of July. Also from Kalso, B. C., collected by
R: B. Currie, western Washington, by H. K. Morrison, Humboldt
County, California, by H. S. Barber, and from several other places in
California and Washington.
152 Annals Entomological Soctety of America [Vol. IT,
Asilus angustipennis n. sp. (Figs. 22 and 23).
Dark species, ground color somewhat obscured by a thin covering
of yellowish dust, wing rather wide, whitish hyaline on the basal part,
brownish beyond the branching of the second and third veins, espec-
ially near the costal border. Length 16-18 mm.
Head rather large, front and face narrow and clothed with golden
dust, facial gibbosity rather prominent but not reaching more than
half way to the antennae, mystax composed of numerous bristles
which are black above and yellow below, beard not abundant, fine
and yellowish white in color; palpi, antennae and proboscis black;
posterior orbits clothed with pale bristles and hairs. Thorax clothed
with yellow dust and black bristles and hairs, the usual black mid-dor-
sal stripe divided before, and some blackish spots on either side;
scutellum with two yellow bristles on its margin and numerous short
light colored hairs on its disk. Wings hyaline at the base but quite
distinctly darkened on the outer half, this darkening is most pro-
nounced along the costal margin before the small cross vein. Legs
black and reddish; base apex and posterior side of each femur, most
of each tibia and bases of the tarsal segments reddish; coxa, trochanter,
most of the anterior side of each femur, apexes of tibiae and tarsal
segments black; many of the bristles of the legs are light, although there
are some blackones. Abdomen clear brown, each segment with anarrow
apical band of pale yellowish preceded on either side by some light
colored bristles. Male genitalia thickened and much wider than the
rest of the abdomen.
A male taken by R. S. Woglum, at Highlands, North Carolina, in
September, 1906, and sent in by Franklin Sherman, Jr. Also a male
from St. Elmo, Virginia, taken by F. C. Pratt, September 13th. The
latter specimen is in the U. S. National Museum.
Asilus latipennis n. sp. (Figs. 24 and 25).
Rather dark in color, hairs and bristles of the body mostly yellow.
Length 15-17 millimeters.
Facial gibbosity rather small, mystax yellowish below, black above,
beard yellowish gray, face above the gibbosity rather narrow and
clothed with golden dust; antennae, palpi and proboscis black, third
antennal segment about as long as the other two, arista about the
length of its segment, bristles and hairs of the occiput mostly pale
yellow. Thorax clothed with yellowish gray dust, with a wide mid-
dorsal stripe and two spots on either side dark brown. Legs black and
yellow with black bristles and yellow hairs, femora largely black in
some specimens, more or less yellowish in others, in all cases observed
the hind femora are yellow at the base; all the tibiae are yellow with
narrow black apexes, base of each first tarsal segment yellowish,
otherwise the tarsi are brown. Wing in the male wide, whitish hya-
line on the basal third, lightly clouded with brownish from thence to
the apex, region of the stigma plainly brown. Wing of the female
hyaline, only obscurely clouded at the apex. Abdomen thinly
clothed with brown dust, posterior margin of each segment yellow,
T9099] ) Robberflies of the Genus Asilus ree
and preceded on either side by several yellow bristles. Male genitalia
rather short, swollen, distinctly wider than the abdomen, shining black
in color; oviduct long and slender, about equal in length to segments
De 0 amd: 7.
A male from Ithaca, New York, collected by Nathan Banks, Aug-
ust 10, 1889, and a male and female from Montgomery, Massachusetts,
collected by Knab, August 24, 1896.
Asilus affinis Williston (Fig. 26).
Mystax of the male bright golden. Legs bright yellow with the
exception of the posterior femora which are black. Length 15 to
16 mm.
Facial gibbosity prominent, mystax composed of abundant golden
hair. Occiput with dark hairs above and pale hairs below. Thorax
black, thinly clothed with dray dust, on the dorsum with numerous
dark more or less bristly hairs; wings hyaline; legs largely yellow, each
middle femur with a black marking above on the basal half, each hind
femur and apex of each hind tibia on the outer side also black. Abdo-
men shining black, apex of each segment with a narrow gray border.
Male forceps wide, excavated on the upper side at the apex.
The male type from which my description and drawing were taken
are in the Francis Huntington Snow Collection at the University of
Kansas. This specimen is from the state of Washington. Another
male taken in the Santa Cruz Mountains of California by Coquillet is
in the U. S. National Museum. :
Asilus flavofemoratus n. n. (Fig. 27).
New name for Asilus flavipes Williston which is preoccupied by
Asilus flavipes Wiedemann from Europe, Syst. Beschr. (1820) II,
320, 28.
Shining black, front and middle femora yellow with the exception
of an elongate black marking above on each, hind femora black, all
the tibiae yellow. Length 12-18 millimeters.
Face and front narrow, facial gibbosity prominent, but not reach-
ing near to the antennae, mystax of the female composed of a few
black hairs above and several white ones below, of the male almost
entirely black, but sometimes partially golden and there may be other
variations. Wings hyaline, very slightly gray at the apex and along
the posterior margin. Each femur of the forward and middle legs
with a black stripe above, hind femora black; all the tibiae yellow, each
posterior black at the apex on the outer side; each metatarsus yellow
with a black apex, other tarsal segments mostly black. Abdomen
shining black, apex of each segment narrowly gray; upper forceps of
the male entire above and of about the same width throughout.
Specimens at hand from Pa., Ohio, N. J., Va., Mass., N. Y., Ills.,
N. C., Md. and Canada. Common in the eastern part of the United
States.
154 Annals Entomological Society of America [Vols LE,
Asilus orphne Walker (Fig. 28).
A black species and in large part shining. Mystax golden, all
the tibiae bright yellow and all the femora black. Wings hyaline.
Length 12-19 “millimeters.
Face narrow, covered with golden dust: facial gibbosity prominent
and extending nearly to the antennae, mystax usually golden in
color but varying to pale yellowish, composed of many slender bristly
hairs; palpi and proboscis black, the former clothed with black hairs;
occiput with black hairs above and pale ones below. Thorax black,
thinly clothed with gray dust, hairs of the sides gray, of the dorsum
mostly black; wings hyaline, very faintly gray at the extreme apex
and along the narrow posterior border. Femora shining black, each
front one “usually yellow at the apex, each tibia and metatarsus yellow,
black at apex, other tarsal segments largely black. Abdomen black,
narrow margin of each segment above gray, clothed with gray hairs
which are most abundant towards the base.
Commonly taken in the territory from Maine to Illinois and from
Canada to North Carolina. Specimens also are at hand from Colorado
and from Montana.
Walker’s description of orphne fits the female of this species well.
Dr. Johannsen agrees with me in identifying auceps. There is no
doubt but that the species under consideration is distinctus and it is
my opinion that this latter is the same as orphne, as is aeneobarbis
which is a name sent by Loew in a letter to Osten Sacken without
description. The name is sufficient to characterize the species
however, when it is known that Loew’s specimens were from the
eastern United States.
Asilus coquillettii n. sp. (Fig. 29).
Black, all the femora black, all the tibiae in large ante yellow. A
prominent crest formed of black bristly hairs extends the whole length
of the thoracic dorsum. In the region covered by this paper no other
species has such a crest. Length 13-16 millimeters.
Facial gibbosity elongate and prominent and reaching nearly to
the antennae. Mystax composed of an abundance of rather fine long
hairs which are all black in the female and black on each side and
otherwise white in the male. Front in both sexes with numerous
long erect black hairs; antennae black, first two segments of each
usually hairy; palpi black with black hairs; lower part of the occiput
with some er ay hairs otherwise with black hairs. Thorax thinly
clothed ht yellowish dust, a crest of long black bristly hairs on the
middle of the dorsum reaches from the anterior part to the scutellum
which also is furnished with numerous similar hairs. Wings hyaline; |
in the male each forward leg has the femur black, the tibia yellow
with a narrow black apex, tarsus yellow with the exception of the
fifth segment and its claws which are black; middle leg the same;
hind leg black with the exception of a little more than basal half of the
tibia which is yellow; in the female the legs are similar but the front
and middle tarsi have the black including the apex of the first segment
1909] Robberflies of the Genus Asilus L565
and nearly all of the other segments. Abdomen shining black with
narrow gray margin to each of the segments.
In some respects this species resembles affinis but the extreme
hairiness of the head and thorax is characteristic, while there is a dis-
tinct difference in the color of the legs and a certain amount of dis-
tinction in the male genitalia.
Males and females collected by Coquillett in Santa Clara County,
California. Types in the United States National Museum.
Asilus terminalis n. sp. (Fig. 30).
Gray dust on the thorax more dense than most species of this
section of the genus, mystax in both sexes black around the outside,
middle part pale yellowish, upper forceps of the male genitalia not
excavated on the dorsal side at the apex. Length 14-17 millimeters.
Facial gibbosity prominent, long but not reaching the antennae,
mystax in both sexes black above and below and with a row of black
bristles on either side, thus the pale yellowish central hairs are entirely
surrounded with black; palpi black with black hairs; antennae black,
first two segments with rather short black hairs; beard white, vestiture
of the upper part of the occiput black. Thorax with a covering of
gray dust which largely obscures the black ground color, dorsum in
front with short and posterior part with longer black hairs; scutellum
with a number of long black bristly hairs on its margin and with num-
erous shorter black hairs on its disk; wings hyaline; front leg in both
sexes with tibia and metatarsus, except the apex of each; and bases
of the other tarsal segments reddish yellow, other parts black but in
some specimens the apex of the femur may be narrowly yellow; middle
leg the same, except there is slightly more reddish yellow at the apex
of each tibia and metatarsus, and the femora are entirely black in all
cases; hind leg black except a little more than the basal half of the
tibia which is reddish yellow. Abdomen black with a narrow gray
posterior margin to each segment and in the female especially each
segment is partially gray on either side. Upper forceps of the male
genitalia not very wide, about the same width throughout, rounded at
the distal end but not excavated at the apex of the dorsal side as in
all other western species of the same section of its genus.
Although this species is not striking in appearance its male geni-
talia mark it as entirely distinct from other known western species.
The male and female from San Antonia Canyon, near Ontario,
California collected July 25, 1907.
Asilus brevicomus n. sp. (Fig. 31).
Black, middle portion of the mystax in the male white, otherwise
black, in the female wholly black, hairs of the whole body decidedly
shorter than in coquilletti1. Length 15-17 millimeters.
Facial gibbosity long and prominent, in the male the mystax
above and below and a row of bristles on either side black, middle por-
tion white, in the female all black; front, and first two segments of the
antennae furnished with short black hairs, occiput with black hairs
above and white ones below. Thorax with short black bristly hairs
156 Annals Entomological Society of America [Vol a,
on the anterior half and longer ones on the posterior half. Wings
hyaline, anal angle in the male with a small white area. Front leg
with the apex of the femur, tibia and metatarsus except the apex of
each, and the bases of the other tarsal segments yellow, otherwise
black; middle leg similar but the femur is entirely black; hind leg
black except three fourths of the tibia which is yellow. Abdomen
shining black with a very narrow posterior margin to each segment
gray. Upper forceps of the male appendages shining black, of nearly
the same width throughout, superior side distinctly excavated at the
apex.
The species is easily separated from affinis by the black forward
and middle femora and from coquillettii by the very much shortened
vestiture especially on the anterior half of the thorax.
Both sexes from Kalso, B. C., sent in by Harvey and Sherman, and
a female from Los Angeles county, California, collected by Coquillett.
Asilus sadytes Walker.
This is the same as Asilus tibialis Macquart, which is preoccupied
by A. tibialis Fabricius, Ent. Syst. (1793) IV, 383, 30; A. tibialis
Fallen from Europe, Dipt. Suec. Asil. (1814) 9, 4; € tibialis Wiede-
mann from Russia, Zool. Mag. (1817) I, 2, 29. There are at least two
other species that have been described under A. tibialis, one from
the Cape by Macquart himself and the other by Gimmerthal from Russia.
Both of these latter have been named since the North American insect.
Black, abdomen with white border to each segment, femora black,
tibiae and metatarsi largely reddish. Male forceps large and dis-
tinctly longer than abdominal segments six, seven and eight. Length
13-17 millimeters.
Face and front clothed with white dust, facial gibbosity promi-
nent, mystax with numerous black hairs above and few white or pale
yellowish ones below, although the distribution of black and light
hairs varies through a series of specimens; antennae black, first two
segments longer than the third which is about equal in length to its
arista. Proboscis and palpi black, the latter with black hair. Beard
white, occipito-orbital bristles black, remainder of the vestiture of the
occiput white. Thorax rather thinly clothed with gray dust which
has a yellowish tinge above, mid-dorsal stripe black; wing hyaline,
apex and posterior margin slightly clouded; coxae colored like the
thorax, femora black, tibiae largely reddish but usually with black
markings on the outside, variable in a series of specimens; each meta-
tarsus reddish with brown apex, other tarsal segments black or brown,
each reddish at the base. Abdomen black with white posterior border
to each segment. Male forceps large and longer than abdominal
segments six, seven and eight.
Specimens are at hand from Ohio, Ind., N. Car., Mass., N. Y. and
Pa. The species is common in sections of Ohio where I have col-
lected in August.
Tgo9] Robberflies of the Genus Asilus sig
Asilus notatus Wiedemann.
Black species. Legs black except the extreme base of each tibiae.
Length 14-18 millimeters.
Face and front clothed with gray dust, facial gibbosity prominent,
mystax with a few black hairs above and numerous pale yellow or
white hairs beneath; beard white; occipito-orbital bristles mostly
_black, other bristles and hairs of the occiput white; proboscis and palpi
black, the latter with black hair. Thorax mostly clothed with gray
dust although the thorax has a yellowish cast; mid-dorsal stripe black,
split anteriorly ; wings hyaline, slightly clouded at the apex and along
the posterior border: the legs are black except the narrow base of
each tibia which is reddish; there is some variation in the legs, the
reddish of the tibiae may be more extensive than indicated and the
metatarsi may be reddish either wholly or in part. It is quite charac-
teristic for this species to have areas of close lying golden pile, especially
on the lower sides of the tibiae and metatarsi. Abdomen black,
each segment narrowly white posteriorly. Upper forceps of the male
genitalia small and short as compared with those of sadytes.
NENG ep Owe Macs Ct, Ils sCanada, Ind. Mos. Kan. Pa.
Me. and Ohio.
Asilus callidus Williston.
Black, each abodminal segment with an apical white band, pre-
apical band of each femur, each tibia and each metatarsus largely
brown. Length 15-18 millimeters.
Facial callosity prominent, mystax black above and pale yellow
or white below. Face and front clothed with gray dust, beard white;
palpi and proboscis black, the former with black hair. Occipito-
orbital bristles variable but always partially black and in some cases
almost wholly black, other hairs and bristles of the occiput pale yel-
low. Thorax clothed with gray dust, mid-dorsal stripe black, narrow-
ly divided anteriorly. Coxae colored like the thorax; femora black
each with a preapical brown band; tibiae brown, each darker on part
of the anterior side and at the apex; tarsal segments largely black,
each brown basally or in case of the metatarsus brown except at the
apex. Wing hyaline, clouded at the apex and along the posterior
border, veins dark colored. Abdomen black, each segment with a
white posterior border; male forceps in large part clothed with pale
yellow hairs, rather suddenly curved at the tip, shorter than abdomi-
nal segments six, seven and eight.
Specimens from British Columbia and Washington. -
Asilus novae-scotiae Macquart.
Dark brown body with plain black femora and yellow tibiae.
Length 14—18 millimeters.
Face and front of ordinary width and clothed with yellow dust,
facial gibbosity rather prominent, mystax composed of black hairs
above and yellowish hairs below, but in some specimens the entire
mystax is composed of light colored hairs; antennae black, third seg-
ment rather narrow and of the same length as its arista; occipito-orbi-
158 Annals Entomological Society of America [Viole iis
tal bristles mostly black above and yellowish at the sides, beard
white. Thorax furnished with yellow dust, mid-dorsal stripe dark,
narrowly divided anteriorly. Legs to the tips of the femora pure black,
tibia yellow with black tips, metatarsi mostly yellow, other tarsal
segments black, each with a narrow yellow base. Abdomen brown
with narrow, light colored posterior margin to each segment. Male
forceps about as long as abdominal segments six, seven and eight, ovi-
duct shining black, first segment equal in length to abdominal seg-
ments six and seven, apical segment slender and style like.
The species is distributed, at least, from North Carolina to Canada
and most specimens have been taken near the Atlantic coast, although
there are records for Ohio.
Asilus antimachus Walker.
A light brown species with clear white mystax and with the legs
almost wholly reddish. Length 16-20 millimeters.
Face and front clothed with gray dust. Facial gibbosity rather
prominent, mystax and beard white; in some specimens there may be
a very few black hairs just above the oral opening and once in a great
while a specimen may be found that has a few short black hairs in the
extreme upper part of the mystax; palpi and proboscis black, the
former with black hair; some of the occipito-orbital bristles may be
black but most of the hairs over the whole occiput are white; antennae
black; first two segments together plainly longer than the third which
is somewhat shorter than its arista. Thorax clothed with gray dust
on the sides, and with yellowish dust above, mid-dorsal stripe and some
spots on either side dark brown. Wings hyaline with slight coloring
at apex and along the posterior border. Coxae colored like the sides
of the thorax, femora largely red in most specimens, but varying from
one having a small elongate black spot on the anterior side to one large-
ly black with a wide preapical band; tibiae yellow, each with the apex
narrowly black or brown, metatarsi yellow, each with a black tip;
other tarsal segments black, some at least with yellow bases. Abdo-
men dull grayish brown, posterior margin of each segment lighter but
the contrast between the segment and its margin not so pronounced
as in novae-scotiae.
Common in various sections of Ohio in late summer. Specimens
also from Indiana, Kansas, Virginia and Missouri. In the locality of
Akron, Ohio, the species is abundant on weeds and in the pastures
during the greater part of August.
Asilus maneei n. sp.
A small black species with black legs, gray stripes and markings on
the thorax and gray posterior borders to the abdominal segments.
Length 10-12 millimeters.
Facial gibbosity rather prominent, mystax composed of a mixture
of gray and black hairs, face and front black but sparsely clothed with
gray dust, beard white, occipito-orbital bristles black, proboscis and
palpi black, the latter with black hair. Thorax black, largely cov-
ered with gray dust, the mid-dorsal stripe black but distinctly divided
1909} Robberflies of the Genus Asilus 159
by a white interval; legs black with not more than a suggestion of red-
dish at the bases of the tibiae, coxae with white hair, femora with some
white hair and black bristles, other parts of the legs with short black
hairs and prominent black bristles; wings hyaline, faintly gray at the
apex and along the posterior border. Abdomen black with rather
long gray hairs especially toward the base, and with the posterior
border of each segment gray. The male appendages shining black,
rather slender, elevated on distal half and about as long as abdominal
segments six, seven and eight together. Oviduct shining black, dis-
tinctly compressed laterally, end lamella free and style lke.
This species must have somewhat the appearance of Asilus anony-
mus Williston, from Mexico, but there are only two bristles on the mar-
gin of the scutellum, the third antennal, segment is of about the same
length as its arista, and there are other differences
A male and female taken in couple at Southern Pines, North Caro-
lina, May 15, 1908, by A. H. Manee for whom the species is named.
Asilus delusus Tucker.
Gray brown. Preapical band and the posterior side of each femur
red. Under side of each front femur with numerous rather long white
hairs. Length 16-18 millimeters.
Mystax with numerous white hairs below and a few black ones
above. Occipito-orbital bristles mostly black above, white on either
side; thorax gray brown, mid-dorsal black stripe well defined, bristles
on the posterior part prominent and black. Wing hyaline, clouded
at the apex and along the posterior border. Front femur with the
anterior side black, preapical band and the posterior side red; tibiae
and tarsi mostly red, each tibia with a stripe on the anterior side and
the apex dark, apex of each tarsal segment brown; middle leg the
same; hind leg similar but the dark coloring of the tibiae more suf-
fused. Abdomen gray brown, narrow posterior margin of each seg-
ment plainly gray.
Specimens from Kansas and Montana, those from Kansas being the
types.
This was described as a variety of annulipes Macquart by Tucker
whose identification of annulipes conforms to specimens called snowit
in this paper, as I have determined by a study of the specimens used
by Tucker. I agree that delusus is related to snowi but has a dis-
tinct general color over the entire body, the posterior side of each
femur red and the third antennal segment shorter than in that species
with the arista two thirds as long as the segment which bears it.
Asilus johnsoni n. sp. (Figs. 32 and 33).
Body bright yellowish, femora black outwardly, otherwise legs
largely red, wing reddish, hyaline along many of the veins. Mid-dorsal
stripe of the thorax prominent, brown and reaching the scutellum in
nearly its entire width. Male genitalia uniformly red. Length 17-21
millimeters.
Facial gibbosity not especially prominent, mystax composed of a
number of rather long pale yellow bristles below and a few shorter
160 Annals Entomological Society of America [Vol. II,
black ones above; antennae black, third segment about as long as the
first two together, arista more than half as long as its segment; occipito-
orbital bristles coarse, black, beard colored like the mystax. Thorax
with a distinct brown mid-dorsal stripe, bristles of the posterior part
of the thorax black and rather prominent; tip of the scutellum with
two black bristles. Front leg with the anterior side of the femur
black, preapical band and the posterior side red; tibia red, marking
on the anterior side and the narrow apex dark; tarsal segments some-
times wholly red but often each is dark at the apex; middle and pos-
terior legs like the anterior ones. Wing reddish, margins of the veins
in large part hyaline. Abdominal segments without distinct lighter
posterior margins; male genitalia shining reddish, upper forceps rather
small, narrow and straight from side view.
Males and females taken at Folson, Delaware County, Pennsyl-
vania, July 7, 1893, by Charles W. Johnson for whom the species is
named.
Asilus snowii n. n.
New name for Asilus annulatus Williston, which is preoccupied by
A. annulatus Fabricius from East Indies, Syst. Ent. 1775, 794, 12;
A. annulatus Macquart from France, Dipt. du Nord 1826, 36, 16.
A rather large dark brown species. Each femur with the excep-
tion of the preapical band, black; tibiae more or less distinctly annu-
late with black; under side of each front femur provided with numer-
ous rather long hairs. Length 15-20 millimeters.
Mystax with black bristles above and white or pale yellowish ones
below; antennae black, third segment long and rather slender, arista
usually not over half the length of its segment. The row of occipito-
orbital bristles mostly black, although in some specimens part of
them are yellow. Thorax clothed with yellowish dust and with the
usual markings and bristles above; wing in large part clouded with
reddish, especially at the apex and along the posterior border, there is
a tendency for many of the veins to be margined with hyaline, even
in the clouded area. Each femur with a preapical reddish band, front
and middle tibiae each with an annulus near the middle and the apex
black, each metatarsus in large part yellow, black at the apex, other
tarsal segments mostly black, usually narrowly yellowish at base;
hind tibiae in large part infuscated, but a narrow basal space is always
yellowish and there appears to be more or less variation in the extent
of the black in a series of specimens. Abdomen clothed with dust so
as to give it a dark brown color, posterior margins of the segments
only slightly lighter.
Specimens are at hand from Canada, N. H., Mo., Penn., N. Y.,
Mass., Ills., Kansas and Ohio.
The type of annulatus is in Philadelphia and Mr. E. T. Cresson, Jr.,
through the kindness of Dr. Skinner, has compared specimens for me.
1909] Robberflies of the Genus Asilus 101
Asilus prairiensis Tucker.
A light brown species, preapical band and the posterior side of
each femur brown. Front femur with close lying hair and several
distinct bristles in a longitudinal row on the under side. Length 14-20
millimeters.
Mystax yellow with the exception of a very few short black hairs
above, occipito-orbital bristles yellow, as are most all of the bristles of
the whole body; third segment of the antenna long, nearly twice the
length of its arista. Anterior femur with a red preapical band and
posterior surface, otherwise black; tibia dark at the middle, especially
on the anterior side and at the apex; metatarsus nearly entirely red,
other tarsal segments mostly black; middle leg like the front one; hind
leg the same except the dark color of the tibia is more suffused. Wing
with the apex and the posterior margin faintly clouded, otherwise
hyaline. In some specimens the hyaline tends to follow the margins
of the veins into the clouded area. Abdomen yellowish brown, the
gray posterior margins of the segments not very plainly marked.
Specimens are at hand from Kansas, Colorado, New Mexico and
Texas:
This species in the southern states appears to occupy about the
same position as paropus at the North. Macquart’s description of
annulipes from Carolina does not apply well to any of the robber-flies
known to me from the northern states, especially is this true of the
words “‘Cuisses fauves, brunatres en dehors,” but does apply to
the present species. These is an older Asilus annulipes by Brullé
from Europe, so the name is not tenable for the American insect.
With a large series of specimens before me showing quite a range in
size and to some extent in coloration, the conclusion is reached that
Tolmerus prairiensis Tucker is the same and being the name next
available becomes the name of the species.
Asilus paropus Walker.
A medium sized species, femora entirely black with the exception of
a reddish preapical ring; Wing reddish although hyaline along the
veins of the disk and the base. Front femur with distinct bristles
below. Length 13-17 millimeters.
Facial gibbosity prominent, mystax composed of numerous bristly
hairs which are mostly pale but there are a very few black ones
above. Antennae black, or at most only narrowly yellow at the apex
of the second and the base of the third segments; occipito-orbital
bristles in large part yellow but variable; in a pair taken in couple,
these bristles are black in the female and yellow in the other sex.
Thorax thinly clothed with yellowish dust and with the usual markings
above, short black hairs anteriorly and long black bristles posteriorly.
Wings distinctly reddish, most of the veins of the disk and toward the
base plainly margined with hyaline. Front leg with the femur, except
a preapical band, middle and apex of the tibia, narrow apex of the
metatarsus and all except the extreme bases of the other tarsal seg-
ments black or infuscated, otherwise yellowish; middle leg like the
162 Annals Entomological Society of America (Vol: iil,
front one; hind leg similar but the tibia except a narrow base may be
wholly infuscated. Specimens are before me in which all the tarsi are
black and the light color of the tibiae is much encroached upon by the
same. Front femur with close lying hairs and a row of distinct bristles
on the under side. Abdomen thinly clothed with yellowish dust, pos-
terior margin to each segment rather obscurely gray.
Specimens are at hand from N. H., Mass., Colo., N. M., Wyo.,
Toronto, Can.-Ct> Nove, Miss ind andl@Ohie:
The species is somewhat variable in size and appearance, but I take
it to be fully distinct. Walker’s description fits quite well here and
prospectus is likely a synonym.
Asilus rubicundus n. sp. (Figs. 20 and 21).
Uniformly reddish, mystax pale yellowish, bristles and hairs on all
parts of the body light colored. Length 12 millimeters,
Face and front rather wide and clothed with yellow dust, facial
gibbosity not very prominent, mystax pale yellowish, beard white,
antennae dark, second segment lighter than the others, third segment
about the length of the other two, arista near the length of its segment.
Occipito-orbital bristles yellow. Thorax yellowish red with a wide
darker mid-dorsal stripe; legs red, outer side of each femur and apex
of each tibia slightly darkened; wings small, hyaline, only a slight
clouding at the apex and along part of the posterior border. Abdo-
men yellow, two or three light bristles on each side before each of the
incisures; genitalia red, upper forceps of the male quite wide at the
tip and evenly rounded.
Specimens of this small and distinct species have been taken at
Onaga, Kansas, by Crevecoeur in June and July, and in southern
Illinois, near Carbondale, also in the same months. The latter speci-
mens are in the Illinois state collection of insects at Urbana.
Asilus mesae Tucker.
Very small species, clothed with pale yellow dust, bristles of the
body and legs, all light colored. Genitaliain both sexesred. Antenna
with a very short arista which is hardly differentiated from its seg-
ment. Length 8-12 millimeters.
Facial gibbosity small, mystax composed of pale bristles, antenna
black, the third segment is characteristic for instead of having the
usual differentiated arista this segment gradually tapers to the apex,
occipito-orbital bristles pale yellow. Ground color of the thorax
hidden by a coating of yellow dust, wings small and hyaline, slightly
clouded at the tip. Femora usually black but variable; tibiae reddish
each darker at the apex, tarsi mostly dark, although each metatarsus
usually is reddish on the basal part. Posterior margin of each abdom-
inal segment narrowly light colored. Genitalia red.
Usually the small size of this species will lead to its recognition,
but it may be confused with rubicundus, from which its antennae will
readily distinguish it.
Specimens are at hand from Colorado, Wyoming and Kansas. The
types are in the Francis Huntington Snow Collection.
1909] Robberflies of the Genus Asilus 163
Asilus erythocnemius n. sp.
_ Dark brown species, each femur except a preapical band black,
other parts of the legs largely red. Third antennal segment rather
wide, arista almost as long as its segment, nearly all the bristles of the
body and legs black. Length 11—14 millimeters.
Facial gibbosity reaching about half way to the antennae, mystax
composed mostly of pale yellow bristles although there are a few short
black ones above, occipito-orbital bristles black, beard white. Thorax
clothed with yellow dust, but with a distinct black mid-dorsal stripe.
Wing hyaline, slightly clouded at the apex and along the posterior
border; femur of each leg black with the exception of a preapical band
which is red; each tibia red, usually with a dark marking near the
middle of the anterior side, apex also dark; metatarsi red, other
segments mostly black. There is a tendency in some specimens
for the hind tibia to be darkened more or less leaving only a narrow
band at the base yellow, especially is this coloring apt to show on
the anterior side. Front femur with a row of rather weak white
bristles on the lower side. Abdomen colored like the thorax, two or
three small bristles on either side before the incisures; from side view
the upper forceps of the male genitalia widest at about two thirds the
length, lower side nearly straight.
Specimens from Mass., N. Y., Ct., N. J., Kan., Mont., Colo., Md.,
Wyo. and Ohio. ~
Asilus astutus Williston (Fig. 35).
Yellowish brown species, form rather robust, femora black anter-
iorly, preapical band and posterior side red. Thorax and legs with
black bristles. Antennal arista about one third as long as the seg-
ment which bears it. Length 15 millimeters.
Facial gibbosity most prominent just above the oral opening, gently
sloping to the level of the face about half way to the antennae. Mys-
tax mostly pale yellow but with several black bristles above, beard
pale yellow; antenna black, third segment distinctly longer than the
first two together, arista about one third as long as the third segment.
Occipito-orbital bristles straw yellow. Thorax clothed with yellow
dust, mid-dorsal stripe dark brown and distinctly split on the anterior
part. Wing in large part clouded, base and disk largely hyaline. Each
femur black anteriorly with a preapical red band, red posteriorly;
front femur with a few black bristles beneath; each tibia dark anter-
iorly with a narrow red base, posterior side red; tarsal segments red,
each more or less darkened apically. Abdomen clothed with yellowish
brown dust, posterior margins of the segments not distinctly lighter;
no bristles before the incisures. In the type male specimen before me
only seven abdominal segments are visible. Genitalia rather small
and red.
The types are in the Francis Huntington Snow Collection and are
labelled as coming from California. There are no other records of the
occurrence of the species known to me,
164 Annals Entomological Soctety of America [Vol aly
If Loew’s names were used this species would come in Antiphrisson
without any question. The form and color taken in connection with
the male genitalia which are figured in the plate, and the absence of
bristles before the margins of the abdominal segments make the species
rather easy of determination.
Asilus californicus n. sp. (Fig. 36).
Dark colored species, wings clear hyaline, legs mostly yellow, pre-
apical bristles of the abdominal segments light colored, upper forceps
of the male genitalia split at the apex. Length 21 millimeters.
Face and front of usual width, facial gibbosity prominent, mystax
with a row of rather fine black bristles around the outside, remainder
composed of coarser white bristles; palp1 black with light hair; antenna
black, third segment about as long as the first two together, arista not
quite as long as its segment; bristles and hairs of the occiput ight with
the exception of a few rather strong bristles just back of the upper
part of each eye. Thorax dark gray, mid-dorsal stripe black but di-
vided by a distinct gray interval. Wings clear hyaline and wider
than usual; front femur black on the anterior side for the basal half
of its length, otherwise yellow; tibia yellow, narrowly black at the
apex; metatarsus yellow, other tarsal segments black; middle leg like
the anterior with the addition or black at the tip of the anterior side
of the femur; posterior leg the same as the middle one except that the
black of the femur takes the form of a stripe not reaching either end.
Abdomen dark gray, posterior margin of each segment only slightly
lighter and preceded by a prominent row of yellow bristles. Male
genitalia shining black with numerous long black bristly hairs especially
beneath; upper forceps rather long, of nearly the same width through-
out, apex split with the lower division the longer and curved inward.
A male of this distinct speeies from Cisco, California, altitude 5,000
feet, taken in June, 1901, and received from Charles W. Johnson.
Asilus sericeus Say.
A large brown species with brown wings and pale brown legs.
Length 20-28 millimeters.
Facial gibbosity with the most prominent point on a level with the
lower edge of the eye, mystax composed entirely of yellow bristles,
palpi black with black hairs, first and second segments of the antennae
yellow, third segment black and decidedly longer than the first two
together, arista about one fourth as long as its segment; bristles and
hairs of the occiput all yellow. Thorax clothed with bright yellow
dust, mid-dorsal stripe dark brown, reaching the scutellum in its full
width; wings brown, veins margined with paler; legs pale brown, each
femur usually with a dark marking on the anterior side. Abdomen
colored like the thorax, no bristles before the incisures.
Generally distributed over the eastern part of North America
from Canada to Texas and at least as far west as Kansas,
1909] Robberflies of the Genus Asilus 165
Asilus midas Brauer.
Entire body and legs velvety black, pulvilli on all the feet pale
brown, hairs and bristles everywhere black except in the male there
is a tuft of white hairs on each cheek below the eye. Wings rich
brownish yellow all over: The species cannot be confused with any
of the others of its subfamily. Brauer called attention to its close
resemblance to a species of Midas from the same locality in Mexico
where the type was taken.
Specimens from New Mexico and from Oak Creek Canyon, Arizona,
taken in August by Dr. F. H. Snow are in the museum of the Univer-
sity of Kansas at Lawrence.
Asilus nitidifacies Hine (Figs. 37 and 38).
Black, wings slightly fumose, face just beneath the antenne shin-
ing black; female with conical oviduct. Length, 12 to 15 mm.
Facial gibbosity rather prominent, and clothed with black bristles
above and white bristles below; face just beneath the antennae shining
black, otherwise clothed with gray dust, which is most pronounced
along the entire margins of the eyes; third segment of the antenna
rather narrow and a little longer than the first two together, arista
only about half as long as its segment, beard white; occipito-orbital
bristles all black. | Ground color of the thorax black, but thinly cov-
ered with gray dust, four to six black bristles on the margin of the
scutellum; wings slightly fumose, almost hyaline on basal parts.
Femora all black, with fine white hair and black bristles; tibiae
and tarsi more or less dark red, approaching black in parts. Hind
tibiae each with three or four black bristles on the front side near the
middle. Abdomen black above, with gray hind borders to the seg-
ments, but not preceded by bristles that differ from those on the other
parts of the abdomen.
A male specimen collected in the Hope Mountains of British Colum-
bia by R. S. Sherman, July 16, 1906, and a female collected on Van-
couver Island, July 2, 1903, by R. V. Harvey. Also specimens in the
United States National Museum from Mt. Hood, Oregon, taken by
H. K. Morrison.
Asilus cacopilogus n. sp.
Gray, wings rather narrow and hyaline. Upper forceps of the male
genitalia not protruding beyond the other part, which is terminated
by a distinct pencil of long white hairs. Length 15-18 millimeters.
Facial gibbosity not prominent, mystax composed of numerous
white bristles, antennae, palpi and broboscis black, palpi and first two
segments of each antenna with black hair, otherwise bristles and hairs
of the whole head white, third antennal segment distinctly shorter
than the first two together, rather wide, arista fully as long as its seg-
ment, front and face clothed with gray dust. Thorax gray, above
with two brown stripes on the anterior part; legs largely black, base
and apex of each femur and basal half or more of each tibia reddish,
the posterior side of each may be partly or entirely reddish, and there
166 Annals Entomological Society of America [Vol. II,
are other variations in the extent of the black and reddish of the legs
in the specimens at hand; claws and pulvilli long and quite slender.
Wings hyaline, posterior branch of the third vein bent slightly forward
at the middle of the length beyond the furcation and then backward
just before the margin of the wing. Abdomen dark, thinly clothed
with gray dust and numerous gray hairs which are longest at the sides.
Ovipositor black with a circlet of spines at the tip; male genitalia rather
short, superior part not protruding beyond the other part which is
tipped with a pencil of long white hairs.
Specimens from Rosser, Texas, taken by F. C. Pratt; Clark County,
Kansas, by F. H. Snow; Anglesif, New Jersey, by H. S. Harbeck,
Havana, Illinois, and from Admore, Oklahoma; Fromont, Nebraska,
and from other places in Texas besides the one mentioned. It appears
to be more eastern in distribution than leucopogon.
This name was carried by some specimens in the Francis Hunting-
ton Snow Collection but I cannot find that any description has ever
been published.
Asilus leucopogon Williston.
Light gray, wings hyaline, scutellum sparsely clothed all over with
rather long white bristles, male genitalia with the superior part pro-
truding at least ha]f its length beyond the inferior part which is with-
out the pencil of hairs present in cacopilogus. Length 15-18 milli-
meters.
Face and front broad, clothed with nearly white dust, facial gib-
bosity most prominent just above the oral opening, gradually sloping
to the level of the face slightly below the insertion of the antennae,
mystax composed entirely of white bristles. Antenna black, third
segment shorter than the first two together, arista about as long as its
segment, palpi and proboscis black, the former with black hair; beard
white, occipito-orbital bristles white or very pale yellowish. Thorax
gray, mid-dorsal brown stripe not conspicuous, wings rather narrow
and hyaline; legs in large part red, each femur on the anterior side with
a black marking which varies in extent in different specimens; each
tibia infuscated toward the tip. Abdomen darkened somewhat and
often with distinct gray posterior margin to each segment. Oviduct
usually shining black, but reddish in part in some specimens, male geni-
talia usually red, superior part extending beyond the inferior part
for about half its length.
Professor Aldrich has kindly sent me a pair of this species from
the material which Williston had when he wrote the original descrip-
tion.
Specimens are at hand from Pierre, South Dakota; from Morton
County, Kansas, and Bill Williams Fork, Arizona, collected by F. H.
Snow. A number of specimens collected by myself near Tombstone,
Arizona, August 5, 1907, are larger than the others and distinctly more
reddish in color but present no structural differences.
1909] Robberflies of the Genus Asilus 167
Asilus arizonensis Williston (Figs. 39, 40 and 41).
Ground color of the body dull black, posterior margin of each
abdominal segment plainly yellowish gray, wings uniformly fumose,
legs reddish. Length 16 to 19 millimeters.
Gibbosity of the face small, clothed with white bristles, sometimes
the two uppermost bristles are black; face and front clothed with gray
dust. Antennae black, or in some cases partially reddish, first seg-
ment of each nearly twice as long as the second, third segment,
without the arista, a little shorter than the first two together, arista
about equal in length to the third segment; bristles of the rear of the
head light colored, although one or two on each side are often black.
Thorax clothed with yellowish gray dust, notum with the usual darker
areas; wings uniformly light smoky brown; legs reddish in general
color, coxae clothed with gray dust and hairs; outer side of each femur,
apex of each tibia and all of each tarsus beyond the apical part of
each metatarsus infuscated. Abdomen dull black above, posterior
border of each segment with a yellowish gray band, sides and venter
gray.
The type of this species is in the Francis Huntington Snow Col-
lection. It is a female and is labelled as coming from Arizona. Be-
sides I have studied a male and two females from southern Arizona
and fifteen males and females taken in the Huachuca Mountains of
Arizona by myself July 28, 1907. Most of the latter specimens were
taken from stones lying at the edge of a small brook where they had
stationed themselves to watch for insects they desired for food.
This is the largest North American species of its group so far
known. The wing in the type is just a shade lighter than in the other
specimens. It is possible that this is due to fading for the specimen
appears as if it is fully developed. The species is very easy to iden-
tify on account of its large size and almost fumose wings.
In a former paper published in the Ohio Naturalist, I identified
the next species as arizonensis, but after studying the type it is evident
that that conclusion was wrong. I did not have specimens of the
present species when I wrote the paper referred to.
Asilus limpidipennis n. sp.
Gray species with hyaline wings and red legs. Body of the male
quite slender, legs distinctly more slender than in arizonensis. Length
17 millimeters.
Gibbosity of the face very small, with a few white bristles; front
and face white; bristles and hairs of the head all white; first segment
of each antenna black, second more or less yellowish and shorter than
the first, third black, exclusive of the arista about as long as the other
two together, arista slender and slightly shorter than the third seg-
ment. Thorax covered with gray dust with a mid-dorsal dark stripe,
narrowly divided before, and two spots on each side, one before and
the other behind the transverse suture. These brown markings are
not so prominent as in some of the other species of the genus. Wings
hyaline, legs largely reddish, coxae colored like the thorax, femora
168 Annals Entomological Society of America [Vol. II,
with more or less black on the outer side, especially on the apical half;
tibiae and metatarsi black at the apexes, other tarsal segments mostly
black. Abdomen dull black, before the incisures with narrow white
bands, in the front margin of which there are on either side in each two
or more bristles.
Male and female taken in south-western Colorado, July 14, 1899,
by E. J. Oslar.
In a former paper published in the Ohio Naturalist I identified this
species as arizonensis but after studying the type of that species have
come to the conclusion that the preceding must bear that name.
Asilus fuscatus n. n.
New name for Philonicus obscurus Hine which is preoccupied by
Asilus obscurus Meigen from Europe, Syst. Beschr. (1820) II, 315, 12.
General color dark with the wings pale fumose all over. The ab-
domen has different shades according to the angle from which it is
viewed, Length 10 to 16 millimeters.
Gibbosity of the face small with a few bristles, part of which are
black and the others white; face with gray dust, front with some
small black bristles and occiput above with a row of black bristles;
antennae black, first segment of each longer than the second, third,
exclusive of the arista, about as long as the other two together; arista
a little shorter than the third segment. Thorax clothed with brown-
ish gray dust, with a mid-dorsal dark brown stripe narrowly divided
before, and on either side two spots of the same color, one before the
transverse suture and the other behind it. Wings uniformly darkened
all over, but it would not be far wrong to say that they are hyaline.
Legs black and reddish; coxae colored like the sides of the thorax,
femora entirely black except that the posterior ones are often narrowly
yellowish at the extreme base of each; tibiae reddish at the base and
black at the apex of each; the extent of the two colors on the tibiae
is somewhat variable, the tendency being for the black to be most
extensive on the outside and the reddish on the inside; feet with the
first two segments of each reddish on basal part, other segments
usually entirely black. Abdomen opaque black, before the incisures
with gray bands, in the front margin of each of which on either side
are two or more distinct bristles.
A male from Kentucky, across the river from Cincinnati, collected
by Charles Dury; two females from Washington, D. C.; males and fe-
males from Riverton, N. J., and Pendleton, N. C., procured from Mr.
Charles W. Johnson; several specimens from New Jersey and Pennsyl-
vania, collected by H. S. Harbeck; also specimens from New York,
Massachusetts and Virginia,
Asilus rufipennis Hine.
General color gray, legs reddish, wings of a uniform reddish color
all over. Length 15 millimeters.
Gibbosity of the face small, furnished with a few bristles which
usually are all white, but in one or two of the species studied there is
now and then a black bristle mixed with the others; face and front
1909] Robberflies of the Genus Asilus 169
with silvery dust; occipito-orbital bristles black; first segment of each
antenna black, second large yellowish and shorter than the first, third
black, not quite as long as the first two together, arista slender, about
the length of the third segment; beard entirely white. Thorax clothed
with gray dust but with a mid-dorsal brown stripe narrowly divided
before and two spots of the same color on either side, one before and
the other behind the transverse suture; coxae gray, usually an elon-
gate blackish spot on the anterior side of each femur and extreme
apexes of all the tibiae and the tarsal segments blackish, otherwise
legs red. | Wings uniformly reddish yellow all over, halteres pale yel-
low. Abdomen opaque black with a narrow gray band in front of each
incisure and in the anterior margin of each of these bands there is on
either side two or more bristles.
Specimens from Douglas County, Kansas, taken in May and June
by E. S. Tucker. Also a specimen from Havana, Illinois, taken June
9, 1905.
Asilus truquii Bellardi (Figs. 42 and 43).
Grayish black, wings pale reddish, thorax with a distinct dark mid-
dorsal stripe; legs slender, femora mostly black anteriorly. Angle at
the middle of the superior side of each part of the male upper forceps
produced inward and backward until it meets and crosses its fellow
of the opposite side. Length 12 to 14 millimeters.
Facial gibbosity very small, mystax composed of a few bristles,
some of the upper ones of which are black and the others white;
antennae black, bristles and hairs of the occiput nearly all gray but
there may be a couple of black bristles behind either eye. Thorax
yellowish gray with a distinct, nearly black mid-dorsal stripe; wings
pale reddish all over; legs slender, femora largely black anteriorly,
reddish posteriorly; each tibia and metatarsus red, dark at the apex;
other tarsal segments in large part black, narrow base of each red.
Abdomen gray black with the posterior border to each segment nar-
rowly gray. Oviduct black, short, conical with four spines at the
tip above.
Specimens from Mexico taken in August. Bellardi, Osten Sacken
and Williston have reported the species from various parts of the same
country.
Asilus tuxpanganus Bellardi. ;
Rather small dark colored species, legs black except a red interval
at the base of each tibia, wings with a slight smoky tinge. Length
11 to 15 millimeters.
Gibbosity of the face small with a few bristles which are black
above and white below; face rather narrow and clothed with white
dust; each antenna black, first two segments with black hairs, second
segment shorter than the first, third slightly shorter than the first two
together. Arista slender and a little shorter than the third segment,
palpi and proboscis black, beard white. Thorax everywhere clothed
with silvery dust, above with two black stripes near the middle of the
170 Annals Entomological Society of America [Vol. II,
notum and two spots of the same color on either side, one before and
the other behind the suture; wings with a slight smoky tinge but this
is so faint that they well may be called hyaline and there is no appar-
ent darkening at the apexes as in many species of the subfamily
Asilinae. Coxae silvery, extreme base of each tibia reddish, other-
wise legs black with black bristles; halteres pale throughout. Abdo-
men shining black above with narrow white bands in front of the incis-
ures; in the anterior border of each white band.there is on each side
two or more medium sized bristles, more prominent in the male than
in the female; venter of the abdomen silvery and with a few white hairs.
Taken at Gualan, Guatemala, January 13th, and at Mazatanango,
Guatemala, February 3, 1905.
EXPLANATION OF THE PLATES.
Fic. 1. Side view of the male genitalia of Asilus gracilis.
Fic. 2. Side view of the female genitalia of A. gracilis.
Fic. 3. Side view of the male genitalia of A. lecythus.
Fic. 4. Side view of the female genitalia of A. lecythus.
Fic. 5. Side view of the male genitalia of A. tenebrosus.
Fic. 6. Side view of the male genitalia of A. avidus.
Fic. 7. Side view of the male genitalia of A. occidentalis.
Fic. 8. Side view of the male genitalia of A. auricomus.
Fic. 9. Dorsal view of the male genitalia of A. auricomus.
Fic. 10. Side view of the male genitalia of A. piceus.
Fic. 11. Dorsal view of the male genitalia of A. piceus.
Fic. 12. Side view of the male genitalia of A. montanus.
Fic. 18. Dorsal view of the male genitalia of A. montanus.
Fic. 14. Side view of the male genitalia of A. albicomus.
Fic. 15. Side view of the male genitalia of A. lepidus.
Fic. 16. Dorsal view of the male genitalia of A. lepidus.
Fic. 17. Side view of the male genitalia of A. willistoni.
Fic. 18. Dorsal view of the male genitalia of A. willistoni.
Fic. 19. Side view of the female genitalia of A. willistoni.
Fic. 20. Side view of the female genitalia of A. rubicundus.
Fic. 21. Side view of the male genitalia of A. rubicundus.
Fic. 22. Side view of the male genitalia of A. angustipennis.
Fic. 28. Dorsal view of the male genitalia of A. angustipennis.
Fic. 24. Side view of the male genitalia of A. latipennis.
Fic. 25. Dorsal view of the male genitalia of A. latipennis.
Fic. 26. Side view of the male genitalia of A. affinis.
Fic. 27. Side view of the male genitalia of A. flavofemoratus.
Fic. 28. Side view of the male genitalia of A. orphne.
Fic. 29. Side view of the male genitalia of A. coquillettii.
Fic. 30. Side view of the male genitalia of A. terminalis.
Fic. 31. Side view,of the male genitalia of A. brevicomus.
Fic. 32. Side view of the male genitalia of A. johnsoni.
Fic. 33. Side view of the female genitalia of A. johnsoni.
Fic. 34. Side view of the male genitalia of A. erythocnemius.
Fic. 35. Side view of the male genitalia of A. astutus.
Fic. 36. .Side view of the male genitalia of A. californicus.
Fic. 37. Side view of the male genitalia of A. nitidifacies.
Fic. 38. Dorsal view of the female genitalia of A. nitidifacies.
Fic. 89. Side view of the male genitalia of A. arizonensis.
Fic. 40. Dorsal view of the male genitalia of A. arizonensis.
Fic. 41. Side view of the female genitalia of A. arizonensis.
Fic. 42. Side view of the male genitalia of A. truquii.
Fic. 43. Dorsal view of the male genitalia of A. truquii.
Pas
Malia i. tot i pO Number 3.
i
oe <BANNA ICS
The Entomological Society of America
SEPTEMBER, 1909
EDITORIAL BOARD,
J. A. COMSTOCK, L. 0. HOWARD,
ITHACA, N, Y. . Par WASHINGTON, D.C.
C.J. S$) BETHUNE, ‘W. M. WHEELER,
GUELPH, ONTARIO, CANADA. ‘ Boston, MAss.
Cc. W. JOHNSON, i P. P. CALVERT;
__ Boston, Mass. PHILADELPHIA, PA.
Wit: KBRLOGE ss J. W. FOLSOM,
STANFORD UNIVv., CAL. URBANA, ILLS.
HERBERT OSBORN, Managing Editer,
COLUMEUS, OHIO.
PUBLISHED’ QUARTERLY BY THE SOCIETY
Entered as ‘second class matter April 11; 1908, at the Post Office at Columbus, Ohio,
undet the Act of Congress of March 3, 1879.
Publications of the Entomological Society of America.
Annals, Vol. I, complete... .2: 6... 02.6.0 -4: Pg SN A CGE Sah SSNS 4
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Annals, Vol. I, Parts 1, 2 and 4, baeh haat a fsastan eta ahs Bi Ray te betta) nite NO ane 1.00
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| . “REPRINTS | es ,
Proceedings of first three nieetings; Coney net By-Laws and . List. of ve
Members... .- EE ADEN GAGNON cetony Ca SAM aM BR EI cea lern eh yea Ry Soke aiiies’3)
WureLer, Wm: M. —Polymorphism Brahe Saat ieee eer ale Pe RGmen
OSBORN, Herperr—The Habits of Insects as a Factor in Classification aed 20
Severin, H..H. anp Severin, H. C: Anatomical and. Histological Studies
of the Female Reproductive Dees of the American- Saw fly; Cimbex .
Americana, Leach. 25th eee = RIS PERL: sie Ree SCS 25
Fxtt, E: P.—Some Problems in Nomenclatnres Bop UE Le Narre Ph Leo Ea
Hammar, A.G.— On the! Neryous System of the der a of Coryaahe cornuta L 28
Bravpiey, J. C.—A Case’ uf Gregatious Sleeping Habits among ‘Aculéate "
Hymenoptera so. 70 Fee tee a Ea ga ae i eae Ea are -10
-Daus,’ J. J—Notes on the Life History of. the Leaiy Dimor as ofthe Box- ’
elder Aphid, Chaitophorus negundinis Thos ... Rais Seite Deh bak LD
HamBieron, J. C.—The Genus Corizus, with a Review ee the North se ;
Middle American Species.:)) 2.0.02 06.0 25 et a ee ee as ta
Giraunr, A. A: — Biological Notes on Colorado Potato. Beetle, . ct) (D5.
Giravit, A; AA? Monographic Catalogue of the) Mymarid Genus Maphus -25
Spverin, H. HH. anp SEVERIN, H. C.—Internal gion of Reproduction of )
Wisie a werfhgy Ao fai Oc a SOE A SR gio ak Stk Ck Ue on ae AS
Smita, C.P.—A Preliminary Study of ine Aranece Theraphosae of California in 75.
Davis, J. J.—Studies on Wphididag, 250 a Bags eae SN cae St N ish Ree tp 4 «20°
Rizzy, W. A—Muscle Attachment of Insects. ..»: PME Tene sDpk oat ee anaes a 15,
NegpuamM, J. G.—Critical Notes on the Classification of the Corduliiae
(Odonata) Sas AR ec Ree Udi aon Re ht keer aid ie St OR RNA Rt tre 4 bp: *
Howarp, L. O.—A Key to the Species of { Pesspaliehs with Table of Hosts >
and Descriptions of Four New Species... ise: diy Ge essere ee SY) ga SN A
Hoop, J. D.—Two New Species of Idolothrips:... 6.20. .2.5.47 Raye : oo) 80.
SAddretet iN iy PAU Yee aes
ANNALS ENTOMOLOGICAL SOCIETY OF AMERICA,
Biological Building, 0. S, U., Columbus, Ohio. «
ANNALS
OF
The Entomological Society of America
Volume II SEPTEMBER, 1909 Number 3
SOME RECORDS OF NORTH AMERICAN GEOPHILIDAE
AND LITHOBIIDAE.
With Description of New Species.
RaLpH V. CHAMBERLIN.
In working over several lots of chilopods from different
parts of the country the members of the Lithobiidae and Geo-
philidae noted below were found. Aside from the addition of
new species, there is reported a number of forms not previously
recognized since the publication of the original descriptions.
A careful examination of a number of species heretofore cata-
logued under Himantarium has shown that these in reality do not
belong to Himantarium as the genus is now conceived, but to
Haplophilus Verhoeff. The new records for Lamyctes fulvicornis
Meinert. definitely establish the range of this form entirely across
the northern portion of the United States. It appears probable
that a second form common in Europe, Schendyla nemorensis
(C. Koch), will prove to have in the United States approximately
the same range as the preceding species, having been previously
reported, however, only from New York State.
1. Schendyla nemorensis (C. Koch).
1837. Geophilus nemorensis C. Koch, Deutschl. Crustacea, etc., Heft 9, Taf. 4.
1866. Schendyla nemorensis Bergsoe 0. Meinert, Nath. Tiddskr. 4 Bd., p. 105.
1880. Schendyla nemorensis, Latzel, Die Myriop. d. Oest.-Ung. Monarchie,
I, p. 198 (which see for the older synonymy.)
1891. Schendyla nemorensis, Cook and Collins, Proc. U.S. N. M., p. 386.
1895. Schendyla nemorensis, Attems, Die Myriop. Stiermarks.
This species, occurring widely over Europe, has previously
been reported in America as occurring at Clyde (Cook and Collins)
and Staten Island (Underwood), New York. Numerous speci-
mens were found by the author under boards and logs in one
particular locality at Provo, Utah, on April 3, t909. These speci-
175
176 Annals Entomological Society of America [Vol. IT,
mens agree closely in mouth parts and other characters with
some from Denmark communicated by Dr. Meinert. Those
examined from Provo have either 39 or 41 pairs of legs.
2. Pectiniunguis montereus Chamberlin.
1904. Pectiniunguis montereus Chamberlin, Proc. Acad. Sci. Phil., p, 653.
Additional specimens, partly grown, from the Monterey Bay
region, agreeing essentially with the types.
3. Pectiniunguis heathii sp. nov.
Ultimate article of antennae in length subequal to the two preced-
ing taken together; a little excavated on exterior side near apex
Cephalic plate eee than wide (as 4.3:3.8); sides widely curved,
but little converging caudad:; anterior margin subtriangular; posterior
margin truncate or weakly curved caudad. Frontal plate not dis-
crete. Prebasal plate not exposed. Basal plate wider than long in
ratio 2:1.
Claws of the prehensorial feet when closed attaining front margin
of head or nearly so; claws and other joints wholly unarmed: prester-
num emarginate mesally in front, the shoes of the emargination meet-
ing at an obtuse angle; wider than long (4.6:3).
Anterior praescuta short, the median and posterior of moderate
length
Each of the first ten ventral scuta with a posterior median process
fitting into a corresponding pit or depression of the succeeding plate.
Ventral pores present on anterior segments in a circular or oval area.
Ventral plates of middle region with a longitudinal median sulcus.
Spiracles all circular, the anterior ones small, the posterior ones
minute.
Ultimate pair of legs each armed with a claw as in montereus, the
claw stout.
Last ventral plate wide, narrowed caudad, each side crenately
excised near its middle; posterior margin widely weakly incurved.
Pleural pores two on ‘each side, moderate i in size and adjacent to the
ventral plate.
Anal pores not detected.
Pairs of legs in female type 45.
Length 22 mm.
Locality—Near Cypress Point, Monterey County, Cal.
Found in an Indian shell mound some distance below surface
A female dug up at Los Angeles and otherwise agreeing with the
type has 53 pairs of legs.
This species is most closely related to the Brazilian P. plusi-
cocitus Attems. It is dedicated to Prof. Harold Heath of Stan-
ford University.
1909] Geophilidae and Lithobiidae 17
~I
4. Escaryus phyllophilus Cook and Collins.
1891. Escaryus phyllophilus, Cook and Collins, Proc. U.S. N. M., p. 392.
This form was found common under leaves, etc., during the
winter months of 1903-04 at Ithaca, N. Y.
5. Haplophilus laticeps (Wood).
1862. Strigamia laticeps, Wood, Journ. Acad. Sci. Phil, V. p. 49.
1865. Strigamia laticeps, Wood, Tr. Am. Phil. Soc., XIII, p. 186.
1886. Himantarium laticeps, Meinert, Proc. Am. Phil. Soc., p. 230.
1903. Himantarium laticeps, Attems, Zool. Jahrb., XIII, p. 176.
Several specimens from Austin, Texas, (March, 1903, Prof.
J. H. Comstock) and from Claremont and Catalina Island, Cali-
fornia (C. F. Baker), agreeing fully with Meinert’s redescription
of Wood's type. Dissection of the mouthparts shows this species
like the two following, to belong to Haplophilus of Verhoetf
6. Haplophilus teniopsis (Wood).
1862. Strigamia teniopsis, Wood, J. Acad. Sci. Phil., V, p. 48.
1865. Strigamia teniopsis, Wood, Trans. Am. Phil. Soc., XIII, p. 185.
1886. Himantarium teniopse, Meinert, Proc. Am. Phil. Soc., a 229,
1889. Himantarium teniopse, Bollman, Proc. U.S. N. M.,
ile
1903. Himantarium teniopse, Attems, Zool. Jahrb., XIII, p. 177.
several specimens from Palo Alto, Cal., (writer, and A. J.
Carlson), where it seems to be common. Meinert has reported
it from San Diego and Bollman from Margarita Island, in the same
State.
7. Haplophilus audacior sp. nov. Pl. X XVI, figs. 1, 2, 3.
Strongly attenuated both cephalad and caudad; smooth.
Antennae short, flattened, broad at base and conspicuously atten-
uated distad. The ultimate article a little shorter than the two pre-
ceding articles together, narrowing distad, a slight depression near
middle of each side.
Cephalic plate clearly wider than long (6:5.3), the anterior margin
widely rounded, not emarginate between the bases of the antennae.
The sides caudad of the anterior rounded portion first nearly straight
or slightly incurved and diverging, then weakly convex and a little
converging to the posterior margin which i is wide and a little excurved.
Frontal plate not discrete. Basal. plate about six times wider than
long..
Labrum free, medianly deeply emarginate, the free margin with
numerous teeth. Mandibles with four pectinate and a single dentate
lamellae, the latter with as many as seven moderately long and api-
cally obtuse teeth; in general structure conforming to type of genus.
First maxillae with inner branch large but not distinct, the outer
branch with second and third segments clearly separated; the coxae
or sternum mostly concealed by those of the second maxillae, each
178 Annals Entomological Society of America [Vol. II,
with a short, inconspicuous palpal process; the second article of the
outer process with a similar palpus which is in a sub-dorsal position
and hence difficult to see. Claw of the palpus of second maxillae
large, smooth.
Claws of the prehensorial feet when closed attaining or nearly at-
taining the front margin of the head; without chitinous lines; unarmed;
front margin of pre-sternum widely though but weakly incurved, not
mesally emarginate, twice as wide as greatest length, or a little wider.
Anterior and posterior dorsal scuta smooth, entirely unfurrowed,
the median scuta somewhat widely depressed each side of middle.
Anterior praescuta very short, becoming caudad gradually longer
until nearly half as long as the scuta.
Ventral plates all smooth and even, unfurrowed and without de-
pressions. Porigerous area transversely elliptical or oblong, in anter-
ior and middle parts of body sharply demarked and somewhat de-
pressed, the lateral portions sometimes more or less bent caudad, in
posterior region becoming more indistinctly defined.
First spiracle large, vertically narrow elliptical, those following cir-
cular, gradually decreasing in size caudad.
The first pair of legs decidedly shorter and more slender than the
second. Ultimate pair of legs six jointed, not crassate and without
a claw.
Last ventral plate narrowed caudad, the sides slightly incurved;
posterior margin nearly straight; anterior margin extending forward
from each side to the middle line, the border being thus sub- triangular:
with a median longitudinal impression. Pleurae with exceedingly
numerous small pores over entire surface, ventral, lateral and dorsal,
some of the ventral pores being covered by the last ventral plate.
Pairs of legs, 97.
Length 102 mm.; greatest width 3 mm.
Locality—Mc Kendrick, Idaho.
8. Geophilus bipuncticeps Wood.
1862. Geophilus bipuncticeps, Wood, J. Acad. Sci. Phil. V, p. 45.
1865. Geophilus bipuncticeps, Wood, Tr. Am. Phil. Soc., XIII, p. 180.
1886. Geophilus georgianus, Meinert, Proc. Am. Phil. Soc. XXIII, p. 218.
1887. Geophilus perforatus McNeil, Proc. U.S. N. M., p. 325.
1893. Geophilus attenuatus, Bollman, Bull. U.S. N. M., 46, p. 148.
Specimens from Columbus, Ohio, and from Washington,
D. C. These have the ventral pores in a transverse band the
anterior margin of which is straight, but the area extends caudad
at middle on some plates, making the posterior portion subtri-
angular. The antennae are long, agreeing essentially with Mei-
nert’s description for geogianus, which is at most a subspecies
of the one under consideration. Bollman, who had McNeil’s
type, regards perforatus as a synonym. Because of a seemingly
strong tendency for the formation of geographical varieties in
1909] Geophilidae and Lithobiidae 179
this form, I reserve judgment as to whether georgianus and _ per-
foratus may not represent such until opportunity shall present
of examining more material or the types.
g. Geophilus attenuatus unaster subsp. nov. Pl. XXV, figs. 4, 5, 6.
Widest anteriorly, strongly narrowing caudad over last fourth of
length.
General color clear yellow; head darker, chestnut over a transverse
area back of frontal region. Antennae uniform yellow. Legs pale
yellow. Venter scarcely paler than dorsum.
Antennae short, strongly attenuated from base distad. Uniformly
covered with fine short hairs. Last article at apex roundly conical,
long, shorter than the two preceding articles together.
Cephalic plate longer than wide (5.2:4); sides straight and subpar-
allel, gradually rounding in to anterior margin, a small but distinct
angular process on each side ectad of base of antenna, widely shallowly
emarginate between bases of antennae; posterior margin slightly
excurved; plate marked with two longitudinal, diverging sulci in front
of posterior margin; with sparse, short hairs, scarcely punctate. Basal
plate essentially free but prebasal plate not showing; not fully two
and a half times wider than long.
Claws of prehensorial feet when closed extending beyond the distal
end of the first antennal article. Claw at base with an acute tooth;
femur near upper inner angle with a stout blunt tooth, each femur
bulging mesad conspicuously adjacent to presternum. Presternum
wider than long in ratio of 5.3:4.7; impressed with a longitudinal
median line; smooth.
Dorsal scuta bisulcate as usual; mostly smooth. Anterior prae-
scuta short, very gradually increasing in length caudad; moderate in
a few segments behind middle, becoming again short posteriorly.
First spiracle large, vertically broadly elliptical, a few following of
the same shape, gradually becoming circular caudad; decreasing in
size from the second caudad, in the posterior region becoming very
small.
Ventral pores in a median longitudinal, lanceolate or ovate area
which extends from the posterior margin cephalad and reaches to or
extends considerably beyond the middle, the area on the anterior seg-
ments more or less sharply demarked but more indefinitely limited
in posterior segments.
Last ventral plate wide, attenuated caudad; each side emarginate
a little below middle, rounded each side of the emargination, the pos-
terior margin being similarly emarginate mesally. Pleurae moderately
enlarged, each with two large pits at the edge of and partly covered
by the last ventral plate.
Anal pores evident.
Pairs of legs 63.
Length 44 mm.; width 1.4 mm.; length of antennae 4 mm.
Locality—Austin, Texas (Prof. J. H. Comstock) ; Louisiana.
180 Annals Entomological Society of America [Vol. II,
10. Geophilus deducens sp. nov. Pl. XXV, figs. 1, 2, 3.
Gradually attenuated cephalad, more abruptly and strongly so
caudad.
Dorsum brown, smoky or blackish along the middle. Venter clear
brown. Head and prehensorial feet pale chestnut, the antennae simi-
lar. Legs pale yellow.
Antennae short. Articles decreasing in length distad, the ultimate
somewhat shorter than the two preceding together.
Cephalic plate longer than wide (4.2:3.7). Posterior margin trun-
cate, anterior margin widely excurved, not mesally emarginate; anter-
ior angles scarcely rounded, the sides caudad of them for ‘about a third
of length straight and diverging, then over middle portion straight
and subparallel, posteriorly rounding mesad to posterior margin.
Roughened posteriorly and laterally. Basal plate about 2.6 times as
wide as the median length. Prebasal plate mesally exposed.
Claws of the prehensorial feet when closed extending beyond front
of head, attaining the middle of the first antennal article. Claw with
a rather pale, short blunt tooth, other joints without teeth. Prester-
num a little wider than long (5:4.8). Presternum coarsely punctate,
the femora nearly smooth or with some finer punctae.
Dorsal scuta bisulcate, some of the anterior ones also with an inner
pair of sulci; roughened. Anterior praescuta short, becoming at and
caudad of middle: moderate- long, not much shortened in posterior seg-
ments.
Spiracles all round; the first large, slightly vertically elongate, the
third being much smaller and the second intermediate in size; caudad
becoming very small.
Anterior ventral scuta with a wide median depression marked with
a longitudinal sulcus which does not extend to either the anterior or
the posterior border, the depression disappearing on posterior segments
and the sulcus becoming longer.
Last ventral plate very wide; the anterior margin on each side
extending forward from side to median line, thus forming a broad
angle; sides nearly straight, strongly converging, the posterior margin
widely excurved. Each pleura with two large pits which are wholly
covered by the last plate.
Anal legs rather short, slender, armed with a long claw.
Anal pores not evident.
Pairs of legs 51.
Length 38 mm. Width 1.3mm. Length of antennae 2.9 mm.
Locality—Sea Cliff, Long Island, New York (N. Banks).
This species is evidently closely related to the attenuatus
group, although clearly distinct.
T9099] Geophilidae and Lith ?btidae 181
11. Geophilus atopleurus sp. nov. Pl. XXIV, figs. 1, 2, 3.
Strongly attenuated cephalad, caudad more abruptly narrowed.
Light, somewhat olivaceous, brown. Head, prehensorial feet and
antennae orange. Legs pale yellow. Venter a little paler than the
dorsum.
Articles of the antennae very gradually decreasing in length distad,
the ultimate shorter than the two preceding together.
Cephalic plate a little longer than broad (12 2:11). Lateral halves
of anterior margin running cephalad. to meet at an angle on the mesal
line. Back of each anterior angle the side is first weakly convex and
the concave at end of frontal suture; back of this the sides are nearly
straight and parallel, caudally rounding in to the wide posterior mar-
gin which is weakly concave. Head marked i in front of the posterior
margin with two diverging sulci. Frontal plate discrete. Prebasal
plate exposed mesally. Basal plate 2.4 times wider than mesal length.
. Claws of prehensorial feet when closed attaining the front margin
of cephalic plate or but little short of it. Tooth on claw very obscure
or obsolete, no trace of one on other joints. Presternum and femora
punctate.
Dorsal scuta sharply impressed with the usual two sulci, which lie
rather close to the median line, some with a more indistinct median
sulcus. Anterior prescuta short, becoming very long at and caudad
of the middle; not much shortened on posterior segments, long.
First spiracle large, circular or a little obliquely elongate; others
circular from the second caudad decreasing gradually but not strongly.
A median process on posterior margin of the second to sixteenth
ventral plates fitting into a corresponding pit or excavation in anterior
margin of succeeding plate. Ventral pores in a transverse band a little
in front of the posterior margin. Sterna with a median longitudinal
sulcus which in anterior segments is short and sharply impressed
but on posterior segments extends over full length of scuta and is
crossed in some by one or more rather indefinite sulci.
Last ventral plate wide; anterior angles rounded; sides somewhat
excurved and strongly converging to the conspicuously rounded pos-
terior margin. Pleurae moderately inflated; each with about eighteen
pores on each side ar ranged along and partly beneath the ventral plate
and along the presterna “and with four or five on dorsal side near and
beneath ine dorsal plate; on the ventral surface caudo-laterad from
the posterior pores a peculiar chitinous mark in a position in some
species occupied by an isolated pore.
Last legs a little enlarged; armed with a large claw.
Anal pores distinct.
Pairs of legs 55 (¢).
Length 39 mm. Width 1.4 mm. Length of antennae 3 mm.
Locality—Raleigh, North Carolina (Brimley, root).
Belonging in the mordax group of species with virginiensis
Boll. and louisianae Brol.
182 Annals Entomological Society of America [Vol. IT,
12. Geophilus rubens Say.
1821. Geophilus rubens, Say, Journ. Acad. Sci. Phil., II, p. 21.
1862. Geophilus cephalicus, Wood, Journ. Acad. Sci. Phil., p. 44.
1862. Geophilus laevis, Wood, Jour. Acad. Sci. Phil., V. p. 44.
1865. Geophilus cephalicus, Wood, Trans. Am. Phil. Soc., XIII, p. 178.
1865. Geophilus laevis, Wood, Ibid., p. 180.
1865. Strigamia rubens, Wood, Ibid., p. 182.
1888. Geophilus rubens, Bollman, Proc. U.S. N. M., XI.
1893. Geophilus rubens, Bollman, Bull. U.S. N. M., 46, p. 148.
1896. Geophilus rubens, Brolemann, Ann. Soc. Ent. de France, p. 53.
Several specimens from Michigan (Ann Arbor, etc.), and two,
representing a variety, from Pacific Grove, Cal.
13. Geophilus smithii Bollman.
1888. Geophilus smithii, Bollman, Proc. U.S. N. M., XI, p. 347.
1888. Geophilus smithii Bollman, Ibid., p. 498.
Four specimens from Washington, D. C., the type locality.
14. Geophilus legiferens sp. nov. Pl. XXIV, figs. 4, 5, 6.
Almost of uniform width over the anterior and middle portions,
attenuated caudad.
Pale orange in general color; head and presternum light chestnut;
dorsum darkened in mesal region; antennae pale orange; legs pale
yellow.
Antennae moderate in length; the articles gradually decreasing in
length distad, the ultimate apically rounded and clearly shorter than
the two preceding taken together.
Cephalic plate longer than wide in the ratio of 4:3. Anterior and
posterior margins substraight or but little excurved, the former slightly
and widely emarginate between bases of antennae. Anterior portion
of sides rounded cephalo-mesad from frontal suture. Sides from a lit-
tle back of ends of frontal suture sub-straight, slightly converging and
posteriorly rounding inward to caudal margin. Front | plate discrete.
Prebasal plate not exposed. Basal plate a little less than three times
wider than the median length.
Claws of prehensorial feet when closed reaching the front margin
of the head. Claws obscurely dentate at base, other joints unarmed.
Anterior prescuta short, becoming moderately long in the middle
region and intermediate in size in the posterior portion.
Anterior spiracle large, vertically widely elliptical, others circular,
the second intermediate in size, gradually decreasing in size from the
third caudad.
First legs shorter and decidedly more slender than the second.
Ultimate legs very long, slender, armed with a large claw.
Sterna uniformly punctate. Ventral pores on anterior sterna few,
in a small median patch just in front of the posterior margin. Anter-
ior sterna mesally widely depressed, marked with a longitudinal
median sulcus.
Last ventral plate long, moderate anteriorly, narrowing decidedly
caudad, the sides weakly excurved. Posterior margin of plate straight.
1909] . Geophilidae and Lithobiidae 18
ios)
Last pleurae each with about twenty-five pores on ventral and lateral
surfaces and a few on dorsal, the pores aggregated chiefly over cephalic
portion, especially in the case of those lateral and dorsal in position.
Two or three pores covered by dorsal plate and four or five by the ven-
tral. The ventral pore most caudad in position is somewhat isolated
from the others but is not enlarged or but slightly so.
Anal pores present.
Pairs of legs 61.
Length 38 mm. Width 1.2 mm. Length of antennae 3 mm.
Locality—-Washington, D. C.
15. Geophilus marginalis Meinert.
1886. Geophilus marginalis, Meinert, Proc. Am. Phil. Soc., X XI, p. 218.
A specimen from Miami, Florida, collected by Prof. J. H. Com-
stock in 1903 agrees closely with the description of Meinert’s indi-
vidual, which likewise came from Florida.
16. Geophilus nasintus sp. nov. Pl. XXIV, figs. 7, 8, 9.
Body gradually attenuated cephalad, more abruptly so caudad.
General color light yellow, inclining to greyish at middle portion
both above and below. Head, presternum and antennae darker, uni-
form.
Antennae long. Articles all long and distad of the second not
much differing in length.
Cephalic plate longer than wide in the ratio of 6:4.8. Posterior
margin subtruncate; anterior angles widely rounded, front margin
rounded forward, mesally not at all emarginate; the sides weakly con-
vex more strongly so posteriorly; posterior margin a little incurved;
head wider behind than in front. Posterior lateral portion of cephalic
plate subdensely punctate somewhat less densely so over median
posterior portion, the punctate rather fine; frontal region almost free
from punetae; frontal plate not discrete. Basal plate three times as
wide as long, rather sparsely finely punctate. Prebasal plate not
exposed.
Claws of prehensorial feet when closed attaining front margin of
cephalic plate. Claw and other joints entirely without teeth. Pre-
sternum nearly smooth.
Dorsal scuta with rather weak lateral sulci, a median longitudinal
sulcus also evident on some; sulci scarcely evident on posterior seg-
ments. Anterior prescuta moderate in length, becoming rather long
in the third quarter of the body and then decreasing in size caudad.
All spiracles round, the first large and the second abruptly smaller.
Ultimate legs in the female long, moderately enlarged, armed with
a claw.
Ventral plates or sterna with a longitudinal sulcus which is deep-
est at middle portion of its length, where on some segments, it is
crossed at right angles by a transverse sulcus. Ventral pores in a
transverse band adjacent to posterior margin.
184 Annals Entomological Society of America [Wolatt
Last ventral plate narrow, the sides converging posteriorly and
the posterior margin straight. Pleural pores situated ventrally
and laterally; a few covered by the last plate and one somewhat
isolated caudo-laterad from the others of the ventral group; sixteen
or eighteen in number.
Anal pores not evident.
Pairs of legs 73.
Length 42 mm. Width 1.1 mm., nearly.
Locality—Not definite, but probably California (southern
portion, if so). One specimen.
17. Geophilus claremontus sp. nov. Plate XXV, figs. 7, 8, 9.
Scarcely attenuated anteriorly; abruptly narrowed caudad.
Cephalic plate longer than wide in ratio of 2.7:2; about equal in
width anteriorly and posteriorly; anterior and posterior angles widely
rounded; posterior margin subtruncate or very weakly excurved; mid-
dle portion of lateral margins straight or but slightly excurved.
Marked with two longitudinal and diverging sulci in front of posterior
margin. Punctate with coarse and fine punctae, the middle portion
of frontal plate free from such punctae or nearly so. Frontal plate
discrete. Portion of the basal plate uncovered by the cephalic 2.6
times wider than long.
Claws of prehensorial feet when closed extending to middle of sec-
ond antennal article. Feet armed on each side with four teeth, that
on the claw longest, truncate; that on the femur broad but low,
rounded, the other two similar but smaller. Presternum wider than
long (4.8:4.5); anterior margin widely emarginate, the emargifation
shallow; marked with a median longitudinal sulcus.
Dorsal scuta strongly bisulcate; punctate. Anterior prescuta
short; prescuta back of the middle becoming moderate and then
decreasing, the ultimate being very short.
First spiracle large, sub-circular, being a little elongate vertically.
Other spiracles circular, the third much smaller than the first and
the second intermediate in size; others gradually decreasing in size
from the third caudad.
First pair of legs decidedly shorter and more slender than the sec-
ond. Ultimate legs slender, armed with a claw.
Ventral pores not detected.
Last ventral plate moderate; narrowing caudad; sides a little ex-
curved as is also the posterior margin. Pleural pores ventral in posi-
tion; the one most posterior in position somewhat isolated from the
others; about fifteen on each side, an inner row of them being covered
by the last plate.
Pairs of legs 65.
Length ad 40 mm. Width ad 1.2 mm.
Locality—Claremont, Cal.
One specimen. Apparently related rather closely to legiferens
sp. Nov.
1909] Geophilidae and Lithobiidae 185
18. Geophilus glaber Bollman.
1887. Geophilus glaber, Bollman, Entom. Americana, II, p. 229.
Pacific Grove and Shasta Springs, Cal. (writer). Common at
Portland, Oregon, (writer) and at Pullman, Washington (Mann).
At Shasta Springs a female was taken in August in a pit on
the under surface of a log with the body coiled about her numer-
ous young.
19. Geophilus occidentalis Meinert.
1886. Geophilus occidentalis, Meinert, Proc., Am. Phil. Soc., p. 220.
One specimen of this species taken at Pacific Grove, Cal., in
June, 1902, and numerous specimens at this place and adjacent
parts in 1909; also at Saucilito (writer).
20. Geophilus regnans Chamberlin.
1904. Geophilus regnans, Chamberlin, Proc. Acad. Sci. Phil., p. 654.
Many additional specimens from southern California (Clare-
mont, Catalina Island, Los Angeles).
21. Geophilus isantus sp. nov. Pl. XXVI, figs. 7, 8, 9.
Ferruginous, becoming brown posteriorly; the head and prehen-
sorial feet darker. Legs yellow. Antennae dark, yellowish distally.
Venter ferruginous anteriorly, becoming yellow caudad. Smooth and
shining.
Body uniform in width anteriorly or nearly so, but attenuated
caudad.
Cephalic plate much longer than wide (4.5:3), wider anteriorly than
behind; anterior and posterior margins truncate, angles rounded, the
gently converging sides nearly straight; anterior margin not mesally
emarginate. Frontal plate not discrete. Basal plate strongly nar-
rowed cephalad, posterior margin 2.7 times as wide as the exposed
portion is long.
Claws of prehensorial feet when closed extending a little beyond
front margin of head. Femur with a blunt black tooth as also each
of two joints distad from it; with a smaller one claw with a tooth at
base which is truncate apically. Presternum acutely emarginate
mesally at anterior border.
Antennae subfiliform, but slightly tapering distad, articles all mod-
erately long, not much slanting toward apex, the ultimate but little
longer than the penult, apically pointed. More than 2.5 times as
long as cephalic plate.
First spiracle large, vertically suboval, the second abruptly smal-
ler, circular, those following gradually and but slightly becoming
smaller.
Anterior ventral plates mesally sulcate. Ventral pores not de-
pected,
186 Annals Entomological Society of America [Vel ais
Last ventral plate longer than wide (nearly as 3.5:2.8), behind
middle bent gently dorsad. Last coxae with four pores arranged at
the corners of a quadrangle and three pores covered by the edge of the
ventral plate on each side.
Anal pores covered.
First pair of legs shorter and a little more slender than the second
Anal legs crassate, seven jointed, the ultimate article unarmed, some-
what tapering distad, whereas the others are more or less obconic.
Pairs of legs 87. Length 40 mm., width 1.1 mm. Antennae 4.4
mm. long.
Locality—Los Angeles, Cal.
22. Linotenia bidens (Wood).
1862. Strigamia bidens, Wood, Jour. Acad. Sci. Phil., V, p. 47.
1865. Strigamia bidens, Wood, Tr. Am. Phil. Soc., XIII, p. 183.
1888. Linotenia bidens, Bollman, Proc. U.S. N. M., XI, p. 34.
1896. Scolioplanes bidens, Brolemann, Ann. Soc. Ent. de France, p. 58, Pl. 6,
figs. 10-13.
One specimen from Raleigh, North Carolina. (Brimley, 1901.)
23. Linotenia fulva (Sager).
1856. Strigamia fulva, Sager, Proc. Phil. Acad. Sci., p. 109.
1862. Strigamia bothriopa, Wood, Jour. Acad. Sci. Phil., V, p. 47.
1865. Strigamia bothriopa, Wood, Tr. Am. Phil. Soc. XIII, p. 183.
1886. Scolioplanes bothriopus, Meinert, Proc. Am. Phil. Soc., X XI, p. 223.
1888. Linotenia fulva, Bollman, Proc. U. S. N. M., XI, p. 341. Ete.
Numerous specimens from the vicinity of lakes Cayuga, Sen-
eca and Keuka, New York (writer, 1903, 1904); Long Island,
New York; Michigan (several specimens).
24. Linotenia chionophila (Wood).
1862. Strigamia chionophila, Wood, Journ. Acad. Sci. Phil., V, p. 50.
1865. Strigamia chionophila, Wood, Tr. Am.: Phil. Soc., XIII, p. 189.
1886. Scolioplanes chionophilus, Meinert, Proc. Am. Phil. Soc., XXI, p. 223.
1888. Linotenia chionophila, Bollman, Proc. U. S. N. M., XI, p. 346.
1896. Scolioplanes chionophilus, Brolemann, Ann. Soc. Ent. de France, p. 60.
Rather common at Ithaca, N. Y., and vicinity (writer); one
specimen from mouth of Mammoth Cave, Ky.; one specimen in
bad condition but apparently this species from Texas.
25. Linotenia levipes (Wood).
1862. Strigamia laevipes, Wood, Journ. Acad. Sci. Phil., V, p. 48.
1865. Strigamia laevipes, Wood, Tr. Am. Phil. Soc., XIII, p. 184.
A number of specimens of this species from Catalina Island,
Cal. Numerous specimens from Los Angeles and Pacific Grove,
occurring in damp soil some distance below the surface.
1g0g| Geophilidae and Lithobudae 187
26. Linotenia rubelliana Chamberiin.
1904. Proc. Acad. Sci. Phil. p. 656.
Additional specimens, adults and partly grown, from the
Monterey Bay region, Cal.; also at Santa Barbara.
27. Lithobius pusio Stuxberg.
1875. Lithobius pusio, Stuxberg, Ofvers. af Kgl. Vet.-Akad. Forhandl., No
2p eOl INO we. ps ole
Claremont, California. Reported originally from San Fran-
CISCO.
28. Lithobius obesus Stuxberg.
1875. Ofvers. af. Kel. Vet._Akad. Frohandl., No. 2, p. 67; No. 3, p. 31.
Specimens from Catalina Island and Claremont, Cal. (C. F.
Baker) ; Laurel Canyon, Los Angeles, and Monterey, Cal. (writer).
29. Lithobius kochii Stuxberg.
1875. Ofvers. af. Kgl. Vet._Akad. Forhandl. No. 2, p. 68; No. 3, p. 30.
1887. Lithobius kochii, Bollman, Proc. U. S. N. M., X, p. 255.
Pacific Grove and Claremont, Cal.; Salina, Colorado (Cock-
erell).
The specimens from Colorado agree with a specimen recorded
from that state by Bollman in having the spines of the first legs
I, I, I aS against o, 1, 1 in those from California. Upon the basis
of Bollman’s statement of this difference Cockerell has proposed for
the Col. form the varietal name coloradensis.
30. Lithobius paradoxus Stuxberg.
1875. Ofvers. af. Kgl. Vet._Akad. Forhandl., No. 2, p. 67; No. 3, p. 31.
A specimen from Claremont, Cal.
31. Lithobius utahensis Chamberlin.
H9OM,” ProciUs Ss. NM, p. 159.
Common at Pacific Grove, and (writer), Shasta Springs, Cal.,
and at Portland, Oregon (writer).
32. Lithobius clavigerens Chamberlin.
1903. Proc. Acad. Sci. Phil., p. 159.
Specimens from Claremont and Catalina Island, Cal. Appar-
ently common along the entire California coast.
33. Lithobius ginampus sp. nov.
Dorsum brown to chestnut, with rather faint darker median stripe;
the head darker; antennae colored like the head at base, distally be-
188 Annals Entomological Society of America [Voli
coming rufous; legs brown, distal joints paler; ventral plates brown,
uniform; presternum reddish brown.
The body conspicuously narrowing from the tenth dorsal plate
forward.
None of the dorsal plates truly produced, with the angles of the
eleventh and thirteenth but weakly extended.
Antennae short; articles twenty-one, these gradually decreasing
in length from the base to the distal one.
Ocelli on each side ten, arranged in three series (144, 3, 2.)
Presternal teeth 2—2.
Coxae of last three pairs of legs armed above, those of the last two
pairs also laterally. Anal legs with the claw unarmed, spines 1, 3, 3, 1.
Penult legs have the claw armed with two spines of which the anterior
one is sub-ventral in position and easily overlooked; spines 1, 3, 3, 2.
Spines of the first legs 1, 3, 2.
Coxal pores round, moderate in size, 3, 4, 4, 3.
Anal legs of the male a little crassate but less so than the penult.
The penult legs have the first tarsal segment strongly inflated and
bearing above at the distal end a conspicuous, short, keel shaped
process.
Length 15.4 mm. Width of tenth plate 2 mm. Length of
antennae 3.6 mm. anal legs short.
Locality—Pullman, Washington (Mann).
The types consist of several males.
34. Lithobius tivius sp. nov.
Dorsum brown, the posterior margins of some scuta bordered with
a somewhat obscure, transverse dark band; head yellowish or yellow-
brown, often of a reddish tinge; antennae dark over the basal portion,
becoming lighter distally, rufous apically; legs yell wish brown, the
posterior pairs darker; venter paler than dorsum, its posterior plates
and also the presternum of rufous caste.
Angles of none of the dorsal plates produced.
Antennae short; articles thirty-one or thirty-two, the first two
articles long, the others, excepting the ultimate, very short.
Ocelli on each side seven or eight, arranged in two series (1+ 3, 3 or
1+4, 3).
Presternal teeth 2—2, moderately large and distinct, darkened.
None of the posterior coxae armed laterally or dorsally. Claw of
each anal leg armed with a single spine; spines 1, 3, 2, 0; moderately
crassate. Penult legs with the claw armed with two spines; spines
1,3, 3, 0. Spines of the first legs OF I
Claw of the female gonopods trilobed, the lobes mostly short and
obtuse, the outside one distinctly smallest; basal spines 2—2, short
and stout. ¢
Length 8 mm. Length of antennae 2.2 mm. Anal legs short.
Locality—Pullman, Washington, and Salt Lake City and
Provo, Utah.
1909] Geophilidae and Lithobiidae 189
This species seems to lie closest to Lithobius exiguus Meinert,
found in New York. In Utah, the new form is commonest under
the bark of decaying logs.
35. Lithobius sokovus sp. nov.
General color of body and legs above and beneath pale yellow to
yellowish white; praesternum and prehensorial feet, and head light
brown: antennae light brown to yellow.
Angles of none of the dorsal plates produced.
‘Head subcordiform, about equal in length and breadth.
Antennae with 19 articles, which are short, the ultimate in length
about equalling the two preceding together.
Ocelli 8 in number, arranged in two series (1+4, 3), decreasing in
size in each series ony most posterior to the anterior one.
Presternal teeth 2—2, very small, brown in color.
Anal legs with claw unarmed; spines 1, 3, 2,0. Penult legs w
claws each armed with a single Spine. Spines of first legs 1, 2, 1.
Coxal pores small, round, Bie Dsl Le
Male: femur of anal legs near middle of dorsal or dorso-mesal
surface with a knot-like elevation, somewhat expanded at distal end
in contact with the tibia. Tibia with a ridge-like elevation extending
for entire length along the dorso-mesal surface, this extended at prox-
mal end into a rounded lobe much as in clavigerens, the lobe bearing a
spine which is curved caudad at its distal end.
Female: claw of gonopods tripartite; basal spines 2-2.
Locality—Las Vegas, Nevada. In loose soil from six inches
to a foot below the surface.
ith
S
36. Lithobius bonvillensis sp. nov.
Dorsum from light to blackish brown, the posterior border of most
of the major plates usually lighter; head paler, brown or light brown,
sometimes with a darker median area caudad of the frontal suture;
antennae brown to blackish, yellowish or rufous distally ; praesternum
and feet concolorous with head; venter light brown to yellowish, the
first and the last plates darkest : legs yellowish, the last few pairs
sometimes darker.
Angles of none of dorsal plates produced.
Head a little wider than long (as 3:2.9).
Antennae short; the first two articles long, others short, closely
crowded; articles 29 in number.
Ocelli 8-9, moderate in size, arranged in two series (1+4, 3 or
Lose)
Praesternal teeth 2—2, small, scarcely darkened.
Posterior coxae unarmed above, laterally or beneath. Anal legs
with the claws each armed with two spines; spines 1, 3, 2,0. Penult
legs with the claws each armed with two spines: spines i aap ap he
Spines of first legs 0, 0, 1.
Igo Annals Entomological Society of America [Vol. I,
Coxal pores small, round, the distal ones in each series largest
3, 4, 4, 4.
Gonopods of female with the claw tripartite, the lobes short and
blunt, the outermost usually less developed than the innermost;
basal spines 3—3, decreasing in size from outermost to innermost on
each side, each spine conically pointed.
Locality.—Lake Point, Utah. Under stones along foothills
37. Lithobius monticola Stuxberg.
5. Ofvers. af Kel. Vet—Akad., No. 2, p. 65; No. 3, p. 30.
Several specimens from Hayward, Cal.
38. Lithobius sierravagus Chamberlin.
1903. Proc. Acad. Sei. Phil., p. 154.
Pullman, Washington, common (Mann).
39. Lithobius xanti (Wood).
1862. Bothropolys xanti, Wood, Journ. Acad. Sci. Phil. V, p. 15.
1865. Bothropolys xanti, Wood, Tr. Am. Phil. Soc., XIII, p. 152.
1875. Lithobius xanti, Stuxberg, Ofvers. af. Kgl. Vet._Akad., No. 3, p. 27.
1887. Lithobius xanti, Bollman, Proc. U. S. N. M., X.
Common at Pacific Grove, Los Angeles, Santa Barbara, San
Bernadino, etc., Cal. (writer).
40. Lithobius multidentatus Newport.
1845. Linn. Soc. Tr. XIX, p. 365.
1862. Bothropolys mobilis, Wood, Journ. Acad. Sci. Phil., V, p. 15.
1865. Bothropolys multidentatus, Wood, Tr. Am. Phil. Soc., XIII, p. 152
Chapel Hill, North Carolina; Virginia; Washington, D. C.;
Michigan; Ithaca, Penn Yan, etc. New York. Common in all
these localities.
41. Lithobius celer Bollman.
1888. L. Entom. Americana, IV.
One specimen, agreeing fully with the description of this species,
from Michigan.
42. Lithobius mordax L. Koch.
1862. Die Myriopodengattung Lithobius, p, 34.
Specimens from Louisiana.
43. Lithopius forficatus (Linnaeus).
1758. Scolopendra forficata, Linnaeus, Syst. Nat., I, p. 638.
1815. Lithobius forficatus, Leach, Tr. Linn. Soc., XI.
1821. Lithobius spinipes, Say, Journ. Acad. Sci. Phil., II, p. 108.
1845. Lithobius americanus, Newport, Tr. Linn. Soc., XIX, p. 365.
1862. Lithobius americanus, Wood, Journ. Acad. Sci. Phil., V, p. 14.
1865. Lithobius americanus, Wood, Tr. Am. Phil. Soc., XIII, p. 148.
1875. Lithobius forficatus, Stuxberg, Ofvers. af Kgl. Vet.-Akad. Forhand1., etc.
1909] Geo philidae and Lithobiidae LQ!
In and about cities and towns of northern Utah (Logan, Og-
den, Salt Lake City, Provo, etc. (writer) ; Pocatello, Idaho (writer).
In Utah I have not found this species away from populated
districts.
44. Lithobius bipunctatus (Wood).
1862. Bothropolys bipunctatus, Wood, Journ. Acad. Sci., Phil., V, p. 16.
1865. Bothropolys bipunctatus Wood, Tr. Am. Phil. Soc. XIII, p. 153.
1875. Lithobius bipunctatus, Stuxberg, Ofvers. af Kgl. Vet.-Akad. Forhandl.
No. 3, p. 30.
1901. Lithobius bipunctatus, Chamberlin, Proc. U.S. N. M., p. 22.
Eastern slope of Wahsatch Mts., Uintah Mts., and Oquirrh
Mts. (Clinton’s Cave), Utah (writer).
45. Lithobius hoples Brolemann.
1896. Ann. Soc. Ant. de France, p. 45.
Three males of this species from Portland, Oregon (writer).
406. Lithobius paucidens a ood.
1862. Journ. Acad. Sci. Phil., 14.
1865. Lithobius paucidens, weed Tr. Am. Phil. Soc., KUMI ps lok
Specimens from Los Angeles, Santa Monica, Laurel Canyon,
1 1G . \
and San Bernadino, Cal.
47. Lithobius harrietae Chamberlin.
1906; Proce AcadyiSe, Phil., p. 3:
Several specimens from Salina, Colorado (Cockerell).
48. Lithobius aztecus Humbert and Saussure.
1869. Rev. et Mag. de Zool., p. 156.
1872. Etudes sur les Myriop., p. 116.
Specimens from Tuxpan and Guadalajara, Mexico.
49. Lamyctes fulvicornis Meinert.
1872. Naturhistorisk. Tiddsskrift, 3rd. R., V, p. 267.
1880. Henicops fulvicornis, Latzel, Die Myriop. I, p. 133. etc.
Lake Point, Utah; Portland, Oregon; Pocatello, Idaho; Las
Vegas, Nevada (writer).
1Q2
Plate XXIV.
Plate X
wo
ot
1
to
Vi.
Annals Entomological Society of America [Vol.
EXPLANATION OF PLATES.
Geophilus atopleurus sp. nov.
Ventral view of anterior portion.
Ventral view of posterior portion.
Dorsal view of anterior portion.
Geophilus legiferens sp. nov.
Ventral view of posterior portion.
Ventral view of anterior portion.
Dorsal view of anterior portion.
Geophilus nasintus sp. nov.
Dorsal view of anterior portion.
Ventral view of anterior portion.
Ventral view of posterior portion.
Geophilus deducems sp. nov.
Ventral view of anterior portion.
Ventral view of posterior portion.
Dorsal view of anterior portion.
Geophilus attenuatus unaster subsp. nov.
Ventral view of posterior portion.
Ventral view of anterior portion.
Dorsal view of anterior portion.
Geophilus claremontus sp. nov.
Dorsal view of anterior portion.
Ventral view of posterior portion.
Ventral view of anterior portion.
Haplophilus audacior sp. nov.
Ventral view of prehensorial feet.
Ventral view of posterior portion.
Dorsal view of anterior portion.
Geophilus isantus sp. nov.
Dorsal view of anterior portion.
Ventral view of posterior portion.
Ventral view of anterior portion.
LT;
ANNALS E. S. A. VOL PLATE XOXLV-.
R. V. Chamberlin.
ANNALS E-S. A.
VoL. II, PLATE XXV.
R. V. Chamberlin.
XXVI.
II, PLAte
Vou.
AS.
Ss.
ANNALS E.
Chamberlin.
1 5
TWO NEW GENERA AND SPECIES OF APHIDIDAE.
By Joun J. Davis,
OFFICE OF THE STATE ENTOMOLOGIST, URBANA, ILLINOIS.
In his “Catalogue des, Aphides “de, Belsique, (+s Dr) a
Schouteden remarks that he has examined specimens of Drepan-
ostphum acertfolia Thomas, and concludes ‘‘qu’il, s’agit d’un
Aphide bien différent de nos Drepanosiphum et qui constituera le
type d'un genre nouveau; la structure des cornicles n’est nullement
celle du genre qui nous occupe.’’ For some time I have consid-
ered the species acerifolit as probably belonging to an unde-
scribed genus. At the suggestion of Mr. H. F. Wilson, of the
U.S. Department of Agriculture, who was also of the opinion that
it belonged to a new genus, I have made a careful study of the
descriptions of Drepanosiphum and its respective species, and
have also had the opportunity of comparing acerifolit with D.
braggu Gil. and D. platanoides Schrank through the courtesies of
Professor C. P. Gillette and Mr. Wilson respectively.
I herewith erect the genus Phymatosiphum to hold a new
species, P. monell1 and that one previously ‘known as Drepan-
ostphum acertfolut Thomas.
Phymatosiphum genus novum.
This genus is at once distinguished from Drepanosiphum, to
which it is closely related, by the cornicles. In Drepanosiphum
they are long and curved, while in Phymatosiphum they are
shorter and decidedly swollen at the base. The stigma of the
former genus, according to the various descriptions and from
examinations by the writer of D. braggi and D. platanoides, is
long and narrow, while in the latter it is shorter and broader, and
has a more distinct whitish central stigmal area.
Description: ' Antenne on frontal tubercles; longer than the body;
the filament of VI long and threadlike, it being longer than any of the
other segments; a single row of large, circular sensoria on the basal
half or two-thirds of segment III. Eyes bright red. Beak rather
short. Wings long and narrow, marginal cell elongated towards the
apex, small dusky patches at the tips of the veins and sometimes a
slight dusky shading along the veins, the stigma brownish excepting a
clear central area (Pl. XXVII, Fig. 1). Fore femur with a prominent
*Mémoires de la Société entomologique de Belgique, tome XII, p. 237, 1906.
196
T9099] Two New Genera and Spectes of Aphididae 107
projection near the distal end (Pl. XXVII, Fig. 2). Two or more
prominent finger-like tubercles on the dorsum of the abdomen.
Cornicles moderately long and dilated at the base. Style small and
globular.
Sporadic in habit. Wingless viviparous adults unknown.
Phymatosiphum monelli, n. sp.
I am indebted to Mr. J. T. Monell for the specimens from
which this species is described, as well as for the very complete
color notes. Mr. Monell has collected this species annually for
the past few years in the months of May and June in St. Louis,
Mo., on the Ohio buckeye (Aesculus glabra). Nearly thirty years
ago he also collected what was perhaps this species, although pos-
itive determination is impossible owing to the loss of the spec-
imens. I quote the following from Monell’s note: ‘‘No. 266x
Oct. 5, 1879: Two dry specimens taken on the underside of
leaves of the buckeye. One winged, general color of dorsum and
antennae light fuscous brown. Short tail yellow. Two tuber-
cles on abdomen. One winged, head and thorax hght fuscous,
the basal half of the abdomen occupied by a dusky transverse
band, the apical half lighter, tail yellow.. Body of winged more
or less pulverulent.”’ To the naked eye it closely resembles
P. acertfolit Thos., but is readily distinguished by the single pair
of dorsal abdominal tubercles, acerzfolit having four pairs of these
tubercles. It lives sporadically on the under surfaces of the
leaves. Mr. Wilson writes that he has found this same species
in Washington, D. C.
Descriptions: Winged viviparous female: To the naked eye this
species resembles P. acertfoli1z. The head is brown with prominent
frontal antennal tubercles (Pl. X XVII, Fig. 3). Antenne brownish
with a dusky to black ring at the distal ends of segments III, IV, V,
and base VI. Segment I large compared with II; III about one and
one-half times as long as 1V; IV and V subequal, V usually being very
slightly the longer; base VI short, less than the length of I; filament VI
longest, being about a half longer than III; total length greater than
twice that of the body. 9 to 15 (usually 11—15) roundish sensoria in
a row on the basal half of III, one large circular sensorium at the
distal end of V, and several surrounding a larger one at distal end of
base VI (Pl. XXVII, Fig. 4). Eyes brilliant red, ocelli three in number
and prominent. Beak rather short, not reaching to the mesocoxe.
Thorax concolorous with the head. Wings hyaline, excepting in the
fore wings a small dusky patch at the end of each vein, at the base of
the stigmal vein, and the stigma which is dark brown with a clear
central spot; venation frail, pale yellowish brown, subcostal dusky
198 Annals Entomological Society of America Vol. LE
yellow; the first and second discoidals branching at a distance of one-
half or slightly more than one-half the distance from the tip of the
wing to where the third branches. (Pl XXXVI) Bic. 1). “fhe arse
discoidal of the hind wing is very faint and invisible in some specimens
in balsam (Pl. XXVII, Fig. 5). In alcoholic and balsam mounted
specimens the legs are brownish and the tarsi black; the femur of each
fore leg bears a prominent projection on the anterior side near the
distal end (Pl. XXVII, Fig. 2). Abdomen covered. with a white
pulverulence which is easily rubbed off, and when it is removed the
color of the dorsum varies from greenish yellow (in recently emerged
adults) to brownish red (in the older ones). On the dorsum is a single
prominent dusky to blackish, double tubercle, apparently borne on
the third abdominal segment (Pl. XXVII, Fig. 6). Cornicles dusky
to black; specimens examined immediately after mounting in balsam
showed the apical half black and the basal half brownish; incrassate
(dilated at the base) and the opening slightly flaring; not reaching to
the tip of the abdomen (Pl. XXVII, Fig. 7). Style pale yellowish and
globular, with a bifid anal plate. (Pl. XXVII, Fig. 8).
Measurements: Length of body, 1.54—1.97, avg., 1.73 mm.;
width, 0.64—0.75, ave., 0.70 mm.; length of wing, 2.82—3.29, aveg.,
3,00; width, 1.04—1.22 avg., 1.10 mm.; total expanse, 6.3—7.0 mm.;
antenna, I, 0.15; II, 0.068; III, 1.00—1.22, avg., 1.15; IV, 0.68—0.86,
avg., 0.78; V, 0.73—0.89, avg., 0.82; VI, base, 0.13; filament, 1.75—
1.9, avg., 1.82; average total, 4.918 mm.; cornicles, avg., 0.32 mm. ;
hind tarsus, 0.14 mm.; style, 0.11 mm.
Pupa:—Light yellowish white to -very pale greenish, with a
longitudinal green stripe on each side of the abdomen. Cornicles
pale greenish.
Described from 6 winged specimens (collected at St. Louis,
Mo., June 30, 1908), on two slides, mounted in balsam from
alcohol, and 5 winged and 5 pupe (collected at St. Louis, Mo.,
May 15, 1908) on two slides, living specimens mounted direct in
balsam, and 9 winged (collected at St. Louis, June 13, 1906), on
one slide, living specimens mounted direct in balsam. All col-
lections made by Mr. J. T. Monell, to whom the writer has the
pleasure of dedicating this species. Two type slides deposited in
the Illinois State Laboratory of Natural History as Accession
No. 40469, one in the U.S. National Museum as Type No. 12630,
and two in the writer’s collection.
Idiopterus genus novum.
Individuals in this genus are small. The head and antennz
similar to those of Phymatostphum. Antenne, set on large con-
spicuous and approximate frontal antennal tubercles (Pl. X XVII,
Fig. 9); longer than the body; segments I and II stout and robust,
T909| Two New Genera and Species of Aphididae 199
the remaining segments setaceous, filament of VI the longest and
decidedly threadlike; circular sensoria in a row on segments III,
IV and V of the winged individuals. Beak comparatively long
and slender. Legs long and frail. Wing venation peculiar and
callipterus-like. Veins with a dark broad border; the stigma
with a light central area similar to that in Phymatosiphum, stig-
mal vein deeply rounded and the fuscous border touching the
discoidal; the third branch of the discoidal is not always distinct,
sometimes it being represented only by a brownish triangular
patch, by a V-shaped dusky patch (Pl. XXVII, Fig. 10), or by
distinct veins, but in any case always very short and at the
extreme tip of wing. Cornicles cylindrical and slightly tapering
from the base, the tips slightly flaring. Style ensiform. The
genus, although bearing some Callipterini characters, appears to
be closer related to the group Aphidini of Mordwilko.
Idiopterus nephrelepidis, n. sp.
This interesting species was first taken in a greenhouse at
Western Springs, Ill., December 2, 1907, on various kinds of
ferns. May 2, 1908, 1t was found quite common and doing some
injury to sword-ferns in a Chicago greenhouse. It is probable
that the species is of tropical origin, as it has only been found on
ferns in greenhouses.
Descriptions: Winged viviparous female: Entire body blackish
or very dark brown. Eyes black. Beak barely reaching the coxe of
the hind pair of legs. Antenne whitish excepting the two basal, the
last segments, and the joints, all of which are blackish; 8—10 circular
sensoria in a row on III, 3—4 in a row on IV, and 2—3 on V;; total
length, exceeding that of the body, segment I more robust and nearly
twice the length of II, III 1-4 to 2-5 Jonger than IV, IV and V sub-
equal, VI (base and filament) subequal to III and IV together, base
VI short and about equal to I in length, the filament very ‘slender and
equal to IV and V together (Pl. XXVII, Fig. 11). Legs whitish
excepting the joints, which are brownish, and the tarsi which are
black. Wings quite characteristic, a conspicuous fuscous border
along each vein, the stigmal vein is subobsolete, as is also the third
branching of the discoidal, which is sometimes almost completely
obsolete. Stigma moderately thick, and with a clear central area.
Other characters of venation as given in the description of the genus.
(Pl. XXVII, Figs. 10 and12). Cornicles cylindrical, sightly imbricate
at base, about twice the length of the style, a a excepting the
basal portion, which is black. (Pl. XXVII, Fig. 13.) Style blackish,
nearly twice as long as the hind tarsus. (PI. SCXVIL. Fig. 14.)
200 Annals Entomological Society of America’ [Vol. II,
Measurements (from specimens mounted in balsam) :—Length of
body, 0.90—1.47, avg., 1.16 mm.; width, 0.40—0.60, avg., 0.50 mm.;
length of wing, 1.78—2.14, avg., 1.98 mm.; width, 0.69—0.83, avg.,
0.76 mm.; avg. expanse, 4.67 mm.; antenna, I, 0.09; II, 0.05;-I1T1;
0.325—.043, avg., 0.3875; IV, 0.21—0.28, avg., 0.245; V, 0.21—0.26,
avg., 0.24; VI, base, 0.09—0.11, avg., 0.10; VI, filament, 0.505—0.57,
avg., 0.54; average total, 1.64 mm.; cornicles, avg., 0.25 mm.; style,
avg., 0.12 mm.; hind tarsus, 0.065 mm.
Described from 14 specimens.
Wingless viviparous female:—Body black with whitish dots scat-
tered over the surface. Eyes black. Antenne pale excepting the
last segment and distal ends of the others; 2 or 3 circular sensoria in a
row on basal half of III; segments about in same proportion as in
winged (Pl. XXVII, Fig. 15). Legs whitish excepting tarsi, which are
fuscous. Cornicles and style as in winged forms.
Measurements (from specimens in “balsam): Length of body,
1.23—1.42, avg., 1.82 mm.; width, 0.65—0.78, avg., 0.73 mm.;
antenna, I, 0.10; II, 0.05; III, 0.836—0.42, avg., 0.38; IV, 0.28—0.26,
avg., 0.24; V, 0.23—0.275, avg., 0.255; VI, base, 0.105; VI, filament,
0.51—0.56, avg., 0.53; average total, 1.66 mm.; cornicles, 0.27 mm.;
style, 0.135 mm.; hind tarsus, 0.08 mm.
Described from 4 specimens.
Two type slides, one deposited in the Illinois State Laboratory
of Natural History as Accessions No. 42533, and the other in the
writer’s collection. One cotype slide in U. S. National Museum
as No. 12629.
EXPLANATION OF PLATE XXVII.
Phymatosiphum monelli n. g. et n. sp., figs. 1-8: Fig. 1, fore wing; Fig. 2,
fore femur, a, basal end; Fig. 3, head; Fig. 4, antenna; Fig. 5, hind wing; Fig. 6,
dorsal abdominal tubercle; Fig. 7, cornicle; Fig. 8, style of winged viviparous
female.
Idiopterus nephrelepidis n. g. et n. sp., figs. 9-15: Fig. 9, head; Fig. 10,
fore wing; Fig. 11, antenna; Fig. 12, hind wing; Fig. 13, cornicle; Fig. 14, style
of winged viviparous female; Fig. 15, antenna of wingless viviparous female.
ANNALS If, S. A. Vou. II, PLate XXVII
John J. Davis
ce Mone ROR cyan ee hea 6 Mambee'
ANNALS
‘The Entomological Society. of America
DECEMBER, 1909
EDITORIAL BOARD.
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STANFORD UNIY., CA. URBANA, ILLS.
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COLUMBUS, OHIO,
_ PUBLISHED QUARTERLY BY THE SOCIETY
Re
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Osporn, Herpert—The Habits. of Insects ds a Factor in Classification..:.:. 20
‘Srverin, H. H. anp Severin, H..C.—Anatomical and. Histological Studies
of the Female Reproductive Organs of the American-Saw fly, Cimbex
Amerieanar beach: ox fos sit open alos a eerie tole se statarars Val Winer dpb Rete 25
ibe A.G.—On the Nervous Syston af the’ Larva of Corydalis aounte A=. 25
Bravery, J. C.—A° Case of Gregarious Sleeping Habits among Aculeate:
Hymenoptera: i oe each ye Bay lant oon Tak Velen cat aie mints pie vie Sta reels -10
Davis, J. J—Notes on the Life History of the Leafy Dimorph of the Box-
elder Aphid, Chaitophorus negundinis Thos... +0... 0.0. eee ee eee e ee 107
HAMBLETON, J. ©The Genus Corizus, with a Reyiew of the North ‘and
Middle-American Species. eles Sag BER acs Ripe ea oie pubinc ete eee
GirAuur, A, A.—Biological Notes on Colorado Potato Beetle... :..: tos ea 25
Girauur, A. A.—A Monographie Catalogue of the Mymarid Genus Alaptus 20.”
Severin, H. H. anv. Severin, H, O—Internal Organs of Réproduction of :
Wie Sa wabhig eho ce eee eed Soa MENS P Oa lola bint wreru tle vis beg vleaee ete boy p © 15
SmiruH, C. P—A Preliminary Study of the Araneze Theraphosae of California +75
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Riuy, W. A.—Muscle Attachment of Insects..... 0.02 ..;505- ec eeeseeeees | 6
Nesepuam, J. G.—Critical Notes on the Classification oF the Corduliinae
(CQmonata).: Sen taserc- cee) Var Res Roe NOMIC ek oie ee SAUDE Aare oI selde LD
Howarp, L. O.—A Key to the Species of Prospaltella with Table of Hosts
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Hoon, J.. D—Two New Species of Idolothrips............5.. 52-4 eee eee by xs
Address
ANNALS ENTOMOLOGICAL SOCIETY OF AMERICA,
Biological Building, O. S. U., Columbus, Ohio
ANNALS
OF
The Entomological Society of America
Volume II DECEMBER, 1909 Number 4
MIMICRY IN THE BUTTERFLIES OF NORTH AMERICA.
By Proressor E. B. Poutton, Oxford, England.
Written from the notes of the Anniversary Address delivered to the Entomolog-
ical Society of America, Baltimore, Thursday, December 31, 1908.
INTRODUCTORY.
Within a few weeks of the hundredth anniversary of Darwin’s
birth, and nearly midway between the fiftieth anniversaries of the
publication of Natural Selection on July 1 last and the Origin of
Species on Noy. 24 next, it seemed to me specially appropriate
to select for this address a subject that is closely associated with
Darwinian teachings. Although he did not publish it during his
lifetime, we now know from.his correspondence that Darwin
independently originated the interpretation of Mimicry which
was afterwards suggested by H. W. Bates. Its development in
the mind of the naturalist of the amazons and the rival theory
afterwards suggested by Fritz Muller were both of them the direct
outcome, in Bates’s case the very speedy outcome, of the Origin.
The deep interest which Darwin took in the hypotheses of both
naturalists is proved by many a letter in his published cor-
respondence. All this forms a peculiarly fascinating chapter of
ancient history,—nevertheless ancient history; but if we desire to
choose a subject because of the light it can throw today and is
certain to throw tomorrow upon evolution and its causes, there is
no study which for promise-as well as performance can be set on
a higher level than Mimicry.
In the course of the following address the word ‘‘Mimicry”’
will be used with the restricted meaning attached to it by A. R.
Wallace. It will be applied solely to the superficial resemblances
between animals, and not to their likeness to vegetable or mineral
surroundings for the purpose of concealment.
203
204 Annals Entomological Society of America [Vol sil:
The study of Mimicry is of the highest value in relation to both
evolution itself and the motive causes of evolution.
Apart from all question of the means by which Mimicry has
been produced, it will be generally admitted that the mimetic
species has in some way evolved a superficial resemblance to the
pattern of one or more species, more or less remote from it in the
scale of classification. Looking on the changes by which the
resemblance has been produced as a piece of evolutionary history,
and, as I have said, disregarding for the moment their causes, we
have one of the very simplest and sharpest pictures of organic
transformation presented for our investigation. An effect—
generally a strongly marked and conspicuous effect—has been
brought about in those elements which make up the superficial
appearance of a species, and this important change is manifestly
in the direction of only a minute fraction of the infinitely complex
organic environment, viz., that fraction contributed by the super-
ficial appearance of one or more very different species, commonly
indeed of but a single one. When, as in North America, a recent
invader becomes the model determining the direction of evolu-
tion in some constituent of the ancient butterfly fauna, the case
becomes especially striking.
The effects produced on the mimic are generally sharper
and more distinct than’ those seen in the concealing resem-
blances to bark, lichen, earth, &c.,—the difference corres-
ponding to the more definite and individual appearance usu-
ally presented by the pattern of the model as compared with such
elements in the vegetable and mineral surroundings. There are
also other important differences. The models of Mimicry are
generally more restricted in their range, and differ more widely
in different areas and in different parts of the same area than the
models of cryptic resemblance. Differences between the local
forms of the same model imply that the mimicked species has
itself been subject to rapid change, while the models of cryptic
resemblance appear by comparison to be stereotyped and _ per-
manent. Furthermore, the models as well as their mimics within
the same area are liable to changes of distribution, whereas the
models of cryptic resemblance are as a rule comparatively fixed.
A mimetic species may often be found passing into an area where
its model exists in a different form or does not exist at all, and
highly instructive conclusions may be drawn from the study of
the corresponding changes.
T909| Mimicry in the Butterflies of North America 205
In accordance with the facts briefly summarized in the above
statements, we find that better and more numerous examples of
rapid recent change are to be found in mimetic patterns than in
those which promote concealment. Not only is this evident
when we trace the geographical changes of model and mimic over
a wide continuous area, but in many cases the same genus
includes both mimetic and non-mimetic species, the latter enabling
us to infer with more or less certainty the ancestral appearance of
the former. The history thus unravelled may often be further
confirmed by a study of the non-mimetic males of mimetic
females.
Many naturalists at the present day incline to return to the
old belief that the history of evolution has been ‘discontinuous,’
proceeding by ‘‘mutations”’ or large and definite steps of change.
The comprehensive and detailed study of Mimicry as a piece of
biological history certainly provides one of the best and safest
means—perhaps the very best—of forming a judgment between
this revived opinion and Darwin's conclusion that, although the
rate of transformation varied greatly and might slow down to
nothing for long periods, the steps of change were small, forming
a gradual and ‘‘continuous”’ transition between the successive
forms in the same evolutionary history.
The study of the causes of Mimicry 1s more difficult than that
of the history of Mimicry, the conclusions far less certain. Nev-
ertheless the evidence at present available yields much support
to the theory of Natural Selection as the motive cause of evolu-
tion. The facts certainly do not point to any other interpreta-
tion. They negative the conclusion that mimetic resemblances
have been produced by the direct action of external forces
(Hypothesis of External Causes) or by variation unguided by
selection (Hypothesis of Internal Causes). Nor do they support
Fritz Muller’s earlier and daring speculation that female prefer-
ences were influenced by the sight of the patterns displayed by
the models (Hypothesis of Sexual Selection). The only hypoth-
eses which are in any way consistent with the body of facts, con-
sidered as a whole, are those which assume that the resemblances
in question have been built up by the selection of variations
beneficial in the struggle for life.
In its concentration on a minute fraction of the total organism
as well as in the rapidity of the results achieved, the operation of
Natural Selection in the production of Mimicry is more than
206 Annals Entomological Society of America [Vol al
ordinarily akin to the methods of Artificial Selection. Indeed a
very fascinating and promising line of investigation in a suitable
locality would be the attempt to initiate or improve a mimetic
likeness by means of Artificial Selection.
Mimetic resemblances are of two kinds, respectively inter-
preted by two well-known hypotheses, both based on the theory
of Natural Selection.
1. Mimicry as interpreted by H. W. Bates is an advan-
tageous deceptive resemblance borne by palatable or harmless
species (the mimics) to others that are unpalatable or otherwise
specially defended (the models). Such resemblance will be
spoken of as Batesian Mimicry, the examples as Batesian mimics,
the interpretation as the Batesian Hypothesis.
2. The resemblances between specially defended species
themselves, although well known to Bates, were not explained by
his hypothesis as he conceived it. He suggested that they were
an expression of the common results produced by forces common
to the environment of the species in question. Such likenesses!
were subsequently interpreted by Fritz Muller as the advantage-
ous adoption of acommon advertisement by specially defended
species, whereby the loss of life incurred during the education of
young inexperienced enemies was contributed between the similar
forms, instead of by each species independently as would have
been the case if they had been dissimilar, and possessed patterns
requiring each a separate education. Such resemblance will be
spoken of as Mullerian Mimicry, the examples as Mullerian
mimics, the interpretation as the Mullerian Hypothesis.
SPECIAL ADVANTAGES OF THE NortH AMERICAN BUTTERFLY
FAUNA FOR THE STUDY OF MIMICRY.
The butterfly fauna of North America affords probably the
best field in which to begin the study of Mimicry,—a subject
which has been shown to possess the most profound significance
in relation to the deepest problems by which the naturalist is
confronted. The examples are sharp and striking, but not too
numerous, and the inquiry can be approached without the con-
fusion and excessive strain on the memory which must inevitably
at first beset the student of Mimicry in the tropics. But outside
the tropics it is also the best field for this study, as will be shown
below.
1It is probable that these were the examples which Fritz Miiller had pre-
viously sought to explain by the theory of Sexual Selection.
1909] Mimicry in the Butterflres of North America 207
The western section of the Palaearctic Region is sharply cut
off by the Sahara from the Ethiopian, and its few examples of
Mimicry are not such as would be likely to awaken the interest
and enthusiasm of the beginner. The eastern Palaearctic section
suffers from the opposite defect. Separated by imperfect bar-
riers from the Oriental Region, its butterfly fauna is complicated
by much invasion of specially protected species from the tropics,
and the examples of Mimicry are too numerous and too little
known. North America occupies a position conveniently inter-
mediate between the two sections of the Palaearctic portion of
the circumpolar land-belt. It has been invaded by models from
the eastern tropics of the Old World and also probably from the
tropics of the New; but the species are few and their effects upon
the indigenous butterflies sharp and distinct. The Mimicry itself
affords striking and remarkable evidence of the lines of migration
followed by some of the intruding models. The ancestral forms
from which the mimics were derived, have nearly always per-
sisted, and enable us to unravel the history of the change, with
exceptional clearness. The examples bear in a most interesting
manner upon the two great hypotheses associated respectively
with the names of H. W. Bates and Fritz Muller. Although the
butterfly fauna is as well known as that of any part of the world,
the mimetic resemblances supply material for a large amount of
much-needed original investigation, inviting the attention of
American naturalists in almost every locality.
THE DaNaINE MopeEts or NortH AMERICA, AND THEIR RELA-
TIONSHIP TO THE SOUTH AMERICAN AND OLD
WoRLD DANAINAE?.
The Danatinae are the most important and most extensively
mimicked of all specially protected butterflies in the Old World
tropics. The Acraeinae, so abundant in Africa, are also greatly
2 The subject of the address from this point onwards is treated in consider-
able detail in the author’s memoir, Mimetic North American species of the Genus
Limenttts (s. 1.) and their models, in Trans. Ent. Soc. Lond., 1908, 447-488. Dr.
Jordan’s later conclusions as to the affinities of Danazda plexippus, added to the
memoir in a terminal note (488) and somewhat at variance with his earlier con-
clusions quoted in the text, are here adopted throughout. <A broader and less
detailed treatment is followed in this address, specialattention being directed to
the numerous points on which further observations are required. Where no
other authority is mentioned, I have followed the synonymy and geographical
distribution of Scudder’s great work, Butterflies of the Eastern United States and
Canada, and, for the Papilionidae, Rothschild and Jordan’s fine monograph
(Nov. Zool., xiii, 1906, 411-752.) I have not, however, followed Scudder in the
general use of Basilarchia as a generic name, because I think that the whole
group of Limenitis, in its widest acceptation, requires revision, and that until
this has been accomplished it is inexpedient to adopt the terminology pro-
posed for a portion of it.
208 Annals Entomological Society of America [Vol. II,
mimicked, but to a far less extent than the comparatively few
species of Danainae found in the same Region,—all belonging to
the section Danaint. The Ethiopian Acraeas in fact supply sev-
eral mimics of the Danaines, but no example of the opposite
relationship is known. In the tropical East, the Acraeznae are
poorly represented, while the Danainae (Danaim, Euploeint,
Hestia, Hamadryas) are dominant in numbers as well as in the
power of influencing the patterns of other butterfly groups. In
both Africa and the East, Mullerian Mimicry is evident between
the different genera and sections of the specially protected groups
themselves.
In the richest and most remarkable -butterfly fauna in the
world, that of South America, the dominant specially protected
group is composed of the /thomiinae, allied to the Danainae, and
called by Bates “‘Danaoid Heliconidae.”’ Next in importance
come the Heliconinae, allied to the Acraeinae, and called by Bates
“‘Acraeoid Heliconidae.’’ Both of these are extensively mim-
icked, especially the [thomiinae: in fact it was the close and obv1-
ous Mimicry of these by certain species of the Heliconinae that
puzzled Bates and ultimately received an interpretation in the
. Millerian Hypothesis. In addition to the above, this rich and
varied Region contains numerous true Acraeimae, mimicked con-
siderably, and a small number of true Danaine species. These
latter, which are of extreme interest, fall into two groups. One of
them, the Lycoraeint, containing the two genera Lycorea and
Ituna, is confined to South America, and bears evident traces of
long residence in the Region. The whole of the species are
mimetic of various dominant Ithomiine genera, while at the
same time some of them appear also to act as models for other
butterflies, in a single case (/tuna phenarete) even for one of the
rarer species (Eutresis imuitatrix) belonging to the Ithominae
themselves. It was the resemblance between the Lycoraeine
genus Ituna and the Ithomiine genus Thyridia that led Fritz
Muller to his hypothesis, and formed the title of the paper in
which he first expounded it. The Lycoraeim are widely different
from any of the Old World Danainae and are sometimes sep-
arated from them as a distinct sub-family. The second group of
Danaines, found in North America, as well as South, belongs to
the Old World section Danain1, and is in every respect strongly
contrasted with the Lycoraeimt. Its species, divided into two
genera Anosia and Tasitia by Moore, are not known to enter into
190 Mimicry in the Butterflies of North America 200
9 y (
mimetic relations with any of the other butterflies of this southern
Region.* Furthermore, they not only belong to a dominant Old
World section of the Danaines, but are even closely allied to par-
ticular species within it. It is probable that there are only two
well-marked species of Danaim on the American Continent, and
that the various forms encountered over this vast area are the
geographical races or sub-species of these two. In north tem-
perate America they are the well-known models for mimicry,—
Anosta plextppus extending far into Canada, and Tasztia berenice
and its form strigosa not ranging beyond the southern States.
In 1897, at the Detroit meeting of the American Association for
the Advancement of Science, I suggested‘ that the Mimicry of
Anosia plexippus by Limenitis (Basilarchia) archippus was
evidence that the model had long resided in North America, and
that we might on this ground alone, even if we had not abundant
positive evidence of its gradually increasing spread in the Old
World during the past half-century, infer that Anosza had reached
Fiji, Australia, Hong-Kong, &c., in comparatively recent times.
This conclusion can hardly be doubted, and the argument might
have been extended to enable us to infer the ancestral line of
migration by which North America itself had been reached by
this form. But in 1897 I followed what appeared to be the gen-
eral view, that, in the New World, the original stream of Danaine
invasion had run from the American tropics northward’, nor did
I observe that the evidence based on the growth of mimetic
resemblance warranted the interesting conclusion that its flow -
had taken the opposite direction, and that the south had been
peopled by way of the north. Accepting this conclusion, the
question arises: Whence came the Danaint of North America?
The answer requires a somewhat careful comparison between the
New and Old World butterflies of this group.
Among the commonest of the Old World Danazint, are certain
species with tawny colouring, a black border, and black white-
barred apex to the fore wing. The under surface is even more
conspicuous than the upper, being brighter in colour and the
black border marked with white in a more striking manner. In
one set of Oriental species, placed by Moore in his genus Salatura,
5 It is possible, however, that there are incipient resemblances to Anosia in
certain S. American Acraeinae.
4 Proc. Am. Assoc. Adu. Sct., 1897, xlvi. 244.
5 Verhandl. d.V. Internat. Zool. Congr. z. Berlin, 1901, Jena, 1902, 171. See
also Essays on Evolution, (1908), 274; also errata.
210 Annals Entomological Society of America [Mola
the veins are heavily marked with black on both surfaces, con-
ferring a very characteristic appearance, especially upon the hind
wing. The other set of species in which the veins are compar-
atively inconspicuous is placed by Moore in Limnas, including
L. chrysippus, perhaps the commonest butterfly in the world,
ranging from the Cape to Hong-Kong and perhaps to Japan. It
is clear, however, that Africa is its ancestral home; for it is there
mimicked far more extensively than in any other country.* In
the Malay Archipelago, both Salatura and Limnas are repre-
sented by various forms, and in some of these the tawny colouring
becomes much darkened. This tendency appears to be more
frequent in Limnas, and when both forms have darkened in the
same island (e. g. Java) it is probable that Limnas has acted as
the model for Salatura. There is a close general resemblance in
colouring and pattern between Salatura of the Old World and
Anosia of the New, as also between Limnas of the Old World and
Tasitia of the New. Furthermore the two New World species
differ from each other in the same points as do those of the Old.
The dark, white-barred apex of the fore wing, so conspicuous in
the Old World forms, is less emphasized in those of the New, being
especially evanescent in Tasitia where, however, traces of the
white markings remain distinct. It is significant, however, that
the black and white apex is also lost in one of the forms of L.
chrystppus, viz., the variety dortppus (= klugw) abundant in
many parts of Africa and also extending by way of Aden and the
west coast of India as far as Ceylon. There is, in fact, much
resemblance between the pattern of dorippus and such a form of
Tasitia as berenice, the likeness being especially apparent in the
indications of the former presence of the white apical bar. In
the forms of Tasitza, as in some of Limnas, the ground-colour
becomes darker and richer—-a development especially well seen
in 7. berentce of Florida. Thus the two chief points in which the
pattern of Tasitia differs from that of typical L. chrysippus, viz.,
the darker, richer ground-colour and the evanescent apical
markings, are both presented by abundant Old World forms of
the latter species. The superficial resemblances between these
Old and New World Danaines are precise and often extend to
minute details. Thus the scent-pouch on the hind wings of the
male, best seen from the under surface, is similar in Salatura and
Anosia, while the resemblance between Limnas and Tasitia in
this respect is even more striking.
§ Proc. Am. Assoc. Adv. Sct.,1.c., 244.
1909] Mimiery in the Butterflies of North America QT
The resemblances above described suggested the investigation
and comparison of structural characters in order still further to
test the relationship between these Old and New World Danaines,
and also the validity of the genera created by Moore.’ Such a
comparison had already been partially made by Rothschild and
Jordan, who in 1903 published the conclusion that Limnas and
Tasit?za cannot be generically separated.’ I therefore wrote to
my friend Dr. Jordan, asking if he would kindly extend his survey
over all the four so-called genera. He found that in Salatura
genutia and Anosia plextppus, having larvae with two pairs of
filaments,® the male genitalia are of the same type; while in
Limnas chrystppus and Tasitia berenice, having larvae with three
pairs of filaments, these genitalia are of a second type. The final
opinion of this distinguished authority on the relationships
between the Rhopalocera, was given in the following words:’
“It appears to be certain that Anosia plextppus does not
stand apart from the others. Therefore, if Tasitza berenice,
Limnas chrystppus and Salatura genutia are placed in one genus,”
plexippus also must be included. I do not think you need
hesitate thus to simplify the classification of these insects.”
I have no hesitation in accepting this advice, and in fusing all
the four genera created by Moore into the single genus Danaida.
Within this genus it has been made evident that the group of
forms ranged around Danaida plextppus is the New World repre-
sentative and close ally of the group of D. genutia; while that of
D. berenice is similarly representative of the group of D. chrysippus
It is interesting to note that both the American Danaidas have
become much larger than the corresponding Old World species,
7 Proc. Zool. Soc. Lond., 1883, 201.
8 Nov. Zool. vol. x, Dec., 1903, 502.
® Dr. Jordan was at first inclined to think that Anosia plextppus should be
separated generically, basing his conclusion in part on the larval characters
(Trans. Ent. Soc. Lond., 1908, 450). A more extended review of the Tring
material pointed in the opposite direction, and Dr. Jordan wrote on December
10, 1908, as follows :—‘I find from our specimens (of preserved larvae) that—
. (1) in Euploea (in the wide sense) there are 4 pairs of filaments, or
three (the 3rd being absent), or two (the 3rd and 4th being absent).
(2) In Danazdae, incl. of Anosia & Limnas, there are 3 pairs (the 3rd of
the 4 pairs of Euploea being absent), or 2 pairs (the 2nd and 3rd being absent).
I find that, for instance, genutia and purpurata have 2 pairs only, like
plextppus. The larva therefore does not furnish anyjargument for sep-
arating plexippus as a genus.’
10 Tn a letter to the author, dated December 15, 1908.
1 Dr. Jordan’s opinion that these three genera should be united is quoted
in Trans. Ent. Soc. Lond., 1908, 450.
212 Annals Entomological Society of America (Vol. i,
and that the most northern forms are larger than the southern in
both hemispheres—the probable result of a slower metamorphosis
in a more temperate climate.
EVIDENCE THAT Danaipa Is AN OLD WoruLpD GENUS THatT HAs
INVADED THE NEw.
_ The suggestion might perhaps be made that the New World
forms of Danaida are the more ancestral, and that those of the
Old World have been derived from them by migration westward.
There is no reason for concluding that the Danaidas of either
geographical area possess a more primitive structure than those
of the other; we are therefore driven to consult other lines of evi-
dence. The following comparisons clearly indicate that Danaida
is an Old World genus which has invaded America at no very
remote period: (1) the far larger number of the Old World
forms and the greater degree of specialization by which some of
them are distinguished; (2) the place of Danaida as one out of a
number of nearly related genera making up the Danainz, a large
and dominant Old World group, per contra its isolated position in
the New World; (3) The highly developed and complex mimetic
relationships of the Old World Danaidas.
This last statement requires some expansion and exemplifi-
cation. Allusion has already been made to the resemblances
which have grown up between different species of Danazda in the
same island,—resemblances in which the forms of chrysippus
appear to act as models. Even more striking is the mimetic
approach of certain Old World Danaidas to species of the other
dominant Oriental section of the Danainae—the Euploeint.
Thus in the Solomons, Danaida (Salatura) insolata is a beautiful
mimic of the dark-white-margined Euploea brenchley1, while in
the same islands Danaida (Salatura) decipiens mimics the dark,
white-spotted Euploea asyllus.’ Finally, and most convincing
as evidence of long residence, are the numbers of mimics which in
the Old World have taken on the superficial appearance of species
of Danaida. In addition to the extraordinary degree to which
the Mimicry of D. chrysippus is carried in Africa, it is mimicked
in the Oriental Region by the females of Hypolimnas musippus
and of Argynnis niphe, and by the males of certain species of
Cethosia. Danaida genutia and the forms related to it are also
2 See J. C. Moulton in Trans. Ent. Soc. Lond., 1908, 603, 604: Pl. XXXIV,
figs. 5, 10.
T909| Mimicry in the Butterflies of North America 21
oN)
mimicked by male Cethosias and extensively by the females of
species of Elymniinae, while incipient Mimicry is seen in the
males of some of them. With the exception of Hypolimnas mist p-
pus, common to both Regions, the Oriental mimics of Danaida do
not approach the degree of resemblance attained by the best
African mimics of D. chrystppus. It has already been pointed
out that the Oriental mimics of this genus are far less numerous
than the African. On the other hand, it is a curious fact that the
only North American mimic of D. plextppus,—Limenitis (Bast-
larchia) archippus—reaches a far higher degree of resemblance
than that attained by any of the characteristically Oriental
mimics of Danazda.
The evidence as a whole enables us to decide that Danazda is
an Old World genus and a compraatively recent intruder into
America, while the perfection of the likeness attained by an
indigenous American mimic proves that, under favourable cir-
cumstances, such resemblances may be rapidly produced. I do
not, of course, mean to imply that the transformation was in
any way sudden, or by other than minute transitional steps. The
evidence for this conclusion will be clearer when some of these
steps have been described in detail. (See pp. 214-17.).
THe LIne oF MIGRATION BY WuHicH DANAIDA ORIGINALLY
ENTERED AMERICA.
There can be little doubt that D. plextppus invaded America
by way of the north, probably following the line of the Aleutian
Islands. In North America it possesses an astonishing distribu-
tion for a member of so tropical a group ranging immensely
further north than any other Danaine in the world. Further-
more, D. genutia, the probable representative of its Old World
ancestor, extends far beyond the tropics into Western and Central
China. A study of the distribution of the Asclepiad food-plants
on the eastern coast of Asia might perhaps throw light on the
problem. D. plexippus was certainly the earlier of the two
invaders of the New World. This is clearly shown by the extent
of its own modification no less than by the changes it has itself
produced. Its immense size, the shape of the hind-wing cell,
and the form of the fore wings indicate that it is far more widely
separated than is D. berenice from the nearest Old World species.
It has furthermore been resident in North America long enough
to effect profound changes in the pattern of an indigenous Nym-
214 Annals Entomological Society of America [Vol. II,
phaline butterfly, rendering it an admirable mimic; whereas
D. berenice, and probably its form strigosa also, have only effected
comparatively slight modifications in the mimetic pattern already
produced under the influence of plexippus (see pp.217-18). It is
impossible to feel equal confidence in suggesting the line by which
the later invasion of the more tropical D. berenice took place; but
it is on the whole probable that it too came by way of the north
during some temporary period of warmth. It is tolerably certain
that it did not invade North America from the south. For
although D. berenice and strigosa have produced—as is shown
above—far less change in the indigenous N. American fauna
than plexippus they have still caused distinct and perfectly effect-
ive modifications in a single species; whereas in South America
their representatives have not been shown to have had any effect
at all. It is probable that both the American Danaidas as they
pressed southward were “‘held up”’ for a considerable time at the
northern borders of the Neotropical Region, unable at first to
penetrate that crowded area. Finally they burst their way
through and are now abundant throughout all the warmer parts
of the Region, the forms of plexippus extending further into the
temperate south, just as in the Northern Hemisphere they range
further north than those of berenice. We are made to realize
the recent date of the invasion of South America when we remem-
ber that nowhere else in the world do Danaine butterflies of equal
abundance ‘‘range through a crowded area without producing
any effect on any member of the Lepidopterous fauna, or without
themselves being affected thereby.”* Abundant wide-ranging
Danaines in the Old World, even when much smaller and with a
less marked appearance, invariably produce some effect, and often
themselves exhibit Mutllerian resemblances.
THE EVvoLuTION OF LIMENITIS (BASILARCHIA) ARCHIPPUS
AS A Mimic OF THE INVADING DANAIDA PLEXIPPUS.
It has already been mentioned that a single species, under-
going corresponding modifications, provides a mimic for each of
the three Danaine models (including strzgosa). We will first
consider the well-known beautiful mimic of D. plextppus; for it
undoubtedly arose earlier than the others.
The abundant Limenitis or Basilarchia archippus is closely
related to the Palaearctic species of Limenitis, a group which
13 Trans. Ent. Soc. Lond., (1908), 452.
1g09| Mimuicry in the Butterflies of North America 21
a |
includes the well-known British “‘White Admiral” (L. sybilla).
The example is unusually instructive, because the non-mimetic
ancestor of the mimic is still very abundant in Canada and the
north-eastern States, and we thus possess the material for recon-
structing the history by which the one form originated from the
other. We know that this ancestor, Limenitis arthemts, has per-
sisted almost unchanged, because of the resemblance between its
pattern and that of the other species of Limenitis (using the name
in the broad sense) from all parts of the cireumpolar land-belt,
including North America itself. The difference between the pat-
tern of the mimic and that of its non-mimetic parent is enormous
—probably as great as that between any two butterflies in the
world; but the steps by which the transition was effected were
long ago suggested by S. H. Scudder,“, and have recently been
worked out in considerable detail by the present writer.’
L. arthemis exhibits the characteristic ‘“‘White Admiral”
pattern—possessing on the upper surface a dark ground-colour
with a broad white band crossing both wings, and white markings
within the apex of the fore wing. Reddish or orange spots be-
tween the white bands and the margin are found in the hind
wings of many individuals, more rarely in the fore wings. These
latter markings are of the utmost importance, for, as Scudder
long ago pointed out (1. c., 714), they undoubtedly provided the
foundation for the change into the mimetic archippus.
A careful comparison between arthemts and archippus reveals
the most conclusive evidence of selection. The one species has
become changed into the other precisely as if an artist were to
paint the pattern of archippus upon the wings of arthemis, retain-
ing unchanged every minute part of the old markings that could
be worked into the new, and obliterating all the rest. Thus,
extending in this direction and wiping out in that, the great trans-
formation has been effected and one of the most beautiful mimics
in the world produced.
The evolution of the mimetic pattern on the under surface has
involved an even more elaborate change than on the upper; but
it is not necessary to repeat here the details which have been only
recently fully described.'® I will, however, allude to the fate of
44 Butterflies of the Eastern United States and Canada, Cambridge, Mass.
(1889), 278, 714.
5 Trans. Ent. Soc. Lond., (1908), 454-460.
16 Trans. Ent. Soc. Lond. (1908), 454-460).
216 Annals Entomological Society of America [Vol. II,
the most conspicuous feature of arthemis, the broad white band
crossing both wings. Save for the traces mentioned below, this
marking has disappeared from both surfaces of the hind wing of
archippus, but its black outer border is retained, and, cutting
across the radiate pattern formed by the strongly blackened veins,
detracts considerably from the mimetic resemblance.'’? On the
under surface distinct traces of the white band may commonly be
seen along the inner edge of the persistent black border. So far
as my experience goes, these traces are only to be found on the
upper surface in the form hulsti (Edw.). The modification of the
same marking in the fore wing is more interesting. Here towards
the costal margin the black outer border is much expanded, in-
vading the white band and cutting off from two to four white
spots from its outer part. While the rest of the band disappears
except on the costa itself, these black-surrounded white spots now
represent the sub-apical pale-spotted black bar of the model. The
new marking is larger and more conspicuous on the under surface,
corresponding with the strong development of white on this
surface of the model. The costal margin of the fore wing of the
latter is streaked with long narrow white markings. In corres-
pondence with this we find, commonly on the under surface, more
rarely on the upper, that the extreme costal end of the white
band is retained, often for the full breadth of the marking, form-
ing a linear streak.
17‘Tn the course of the address on December 31, 1908, I remarked that if we
could revisit the earth in a few hundred years, we might expect to find that this
black line had disappeared from the hind wing, and the mimetic resemblance
correspondingly heightened. At the conclusion, Mr. John H. Cook, of Albany,
N. ¥., informed me that he had discovered near his home many indviudals in
which the black line was wanting from the upper surface. A few days later he
very kindly sent me a record of his observations, of which an abstract is printed
as a note at the end of this address (see pp. 241-42). The study of Mr. Cook’s
facts shows that near the city of Albany not only did the stripeless variety occur
commonly (1 in 14), during the three seasons in which the observations were
conducted, but also transitional forms with more or less broken stripes were far
commoner than the normal archippus (18 to 1). The fact that entirely stripe-
less individuals were invariably males is contrary to the rule that mimetic
resemblance tends to develop more rapidly and fully in the other sex. But in
this species I have observed another point in which the female tends to be more
ancestral than the male, viz., the more frequent and complete development of
the white spot in the cell of the fore-wing upper surface (a common feature of
Limenitis, although now generally absent from L. arthemts).
Mr. Cook’s observations show that a single marking—and one so simple
that we might have expected it to act as a unit, so small a traction of the pattern
that we could hardly speak of its sudden disappearance as an example of ‘‘dis-
continuous’’ evolution—that even this behaves differently on the two surfaces
of the wing, while the individuals from which it has disappeared are immensely
outnumbered by those in which it is transitional.
1909 | Mimicry in the Butterflies of North America 207
I have dwelt upon the changes undergone by the white band
as an example of the way in which the new markings have been
carved out of the old. The changes in the elaborate marginal
pattern would have been equally convincing as evidence for a
gradual and “‘continuous”’ transformation.
Tue MopiFICATION OF THE LIMENITIS MiImMiIc oF DANAIDA
PLEXIPPUS INTO A Mimic oF D. BERENICE IN FLORIDA.
Danaida plexippus occurs together with D. berenice in Florida,
but the latter far outnumbers the former, and the modification of
Limenttis archippus into the form floridensis, Strecker (=eros,
Edw.) is probably entirely due to the predominance of one model
over the other. Data for determining the exact proportions in
various localities would be of high interest. There is no reason
for believing that berenice is in any way more or less distasteful
than plexippus, but its abundance makes it a more conspicuous
feature in the environment.
It is evident that the change has been of the kind expressed
in the above heading; for, ashas been already implied in pp. 213-14,
traces of the former Mimicry of plexippus persist in floridensis
and tend to detract from the resemblance more recently developed.
This is especially the case with the conspicuously blackened veins
of archippus, which are so important a feature in the likeness to
plexippus. These, although obscured by the general darkening,
are still recognizable in floridensis, diminishing its resemblance
to berenice on the upper surface of both wings and on the under
surface of the fore wing. Inasmuch as the details have been
recently published elsewhere,'* I will only dwell on one further
point in the resemblance of floridensts to berenice—and that be-
cause the extensive observation of large numbers of specimens is
greatly needed. I spoke on pp. 215-16 of the persistent traces of
the white band on the hind-wing under surface in many indivi-
duals of L. archtppus. These are ancestral features, diminishing
the mimetic resemblance to D. plexippus. But in D. berenice
there are conspicuous white spots towards the centre of the hind-
wing under surface, and these, at any rate upon the wing, would
bear some resemblance to the ancestral spots of the Limenitis
mimic. Now in my very limited experience of floridensts these
spots were sometimes exceptionally developed and, outlined with
black on their inner edges, were certainly far more distinct and
18 Trans. Ent. Soc. Lond. (1908), 460, 461. See also Scudder, 1. c. 718.
218 Annals Entomological Society of America [Vol. IT,
conspicuous than in L. archippus. The appearances I witnessed
suggested the possibility of the recall of a vanishing feature in
consequence of selection based on a likeness to certain white
spots present in the new model (berenice) but absent from the
old (plexippus). But many hundreds of specimens from differ-
ent localities scattered over the total area of distribution require
to be examined from this point of view. An even more interest-
ing inquiry would be to trace the range of the floridensis form
northward and determine the relationship of its limits to the zone
in which berenice becomes scarce and disappears, and above all
to ascertain whether floridensis on the borders of its range inter-
breeds with archippus and how far transitional varieties occur.
Interbreeding between the two forms, if possible, would be of
extraordinary interest. It is also of importance to ascertain
precisely how far the one form penetrates the area of the other.
Scudder indeed states that floridensis ranges into the Mississippi
Valley and Dakota, far beyond the limits of Danaida berenice.
It would be deeply interesting to make an exact comparison
between such specimens and those from Florida, and also to
ascertain the proportion which they bear to typical archippus.
By far the most important feature in the evolution of floridensis
is the general darkening of the ground-colour, and the material
for such a transformation certainly exists freely in archippus, for
the shade of brown varies immensely and may often be seen of as
dark a tint as in floridensis, but not in my experience of precisely
the same shade. The proportion of such dark forms in various
parts of the immense range of archippus would be another inter-
esting inquiry.
Tue MopiIFICATION OF THE LIMENITIS Mimic oF DANAIDA
PLEXIPPUS INTO A~ MIMIC OF THE STRIGOSA FORM OF
D. BERENICE IN ARIZONA.
The differences between L. archippus and the form hulsti
(Edw.) are more striking than those which distinguish floridensts
from the former. The upper surface of the hind wing of Mulsti
retains or more probably has recalled distinct traces of the white
band, although the black stripe is evanescent. It is probable that
upon the wing, these vestigial white markings produce a general
likeness to the pale-streaked hind-wing upper surface of strigosa.
Other points in which hulst: differs from archippus and approaches
strigosa are the reduction of black and the general appearance of
1909] Muimuicry in the Butterflies of North America 219
the white spots in the subapical region of the fore wing, and the
dull tint of the ground-colour. I have had hardly any exper-
ience of this interesting form and owe the above details to Dr.
W. J. Holland’s figure and description.'® It is obvious that all
the investigations suggested in the case of floridensts are, mutatis
mutandis, equally available and equally important in the form
hulstt.
The geographical distribution of hulstt strongly supports the
conclusion that it was derived from archippus and not 1mme-
diately from an arthemts-like ancestor. I have not yet had the
opportunity of ascertaining whether this hypothesis is supported
by evidence derived from a careful study of the pattern.
It is deeply interesting to observe that the same Limenitis
arthemts-like species, from which .archippus, floridensis, and
hulsti—mimics respectively of the three Danaidas, plexippus,
berentce and strigosa—have been directly or indirectly evolved,
has also given rise to L. astyanax (ursula), the mimic of a Papil-
ionine model. Evidence in favour of the comparatively recent
origin of these mimicking forms is to be found in the well-sup-
ported facts which indicate that astyanax still interbreeds with
arthemis along their geographical overlap, and that it may even
occasionally pair with the sister species archippus.*°
The earlier stages of archippus and astyanax are, according to
Scudder (1. c. 254, 255), with difficulty distinguished from those
of arthemis, but astyanax presents the closer likeness of the two;
a'fact which, together with those referred to in the last para-
graph, points to the conclusion that it arose even more recently
than archippus,
The further consideration of astyanax is best deferred until
some account has been given of the Papilionine models, and until
certain general conclusions have been discussed in the following
section.
BEARING UPON THEORIES OF MIMICRY OF THE TRANSFOR-
MATION WROUGHT BY THE INVADING DANAIDAS.
It has been shown that the Danaine models invaded America
from the Old World tropics, probably following a northward route.
Their patterns are but little changed in the new surroundings,
and they still keep the characteristic appearance of Old World
19 Butterfly Book, 84, 185, Pl. vii. f. 5. Dr. Holland fully recognizes the
mimetic significance of the pattern and colouring of hulstt.
20 Scudder, 1. c. 283, 289. Trans. Ent. Soc. Lond. (1908), 473, 474.
20 Annals Entomological Society of America [ Vol. 1f,
bo
Danaidas. Furthermore, such changes as have taken place in
the older invader, D. plextppus, during its residence in the New
World, are also retained in those colonies which, during the half
century, have been re-establishing themselves in the Old World.
These facts support Darwin’s conclusion that the physico-
chemical influences of soil, climate, etc., are of comparatively
slight importance, a conclusion which made him feel ‘‘inclined
to swear at the North Pole, and * * * to speak disrespectfully of
the Equator.’’”?
The mimics on the other hand are derived from characteristic
and ancient inhabitants of the northern land-belt. If, as the fol-
lowers of the theory of External Causes (see p. 205) maintain,
species are the expression of the physical and chemical forces of
the environment, then the Danaidas express the Old World
tropics and the species of Limenitis the northern land-belt. We
might expect on this theory that the Danaidas, when they in-
vaded the northern zone, might come to resemble the Limenztts;
but the transformation that has actually occurred is entirely
inconsistent with any such hypothesis. Although the Danaidas
have undergone no important change in the new environment,
their presence has entirely transformed and brought into a close
superficial resemblance to themselves the descendants of a mem-
ber of an ancient group. Such a fact is inconsistent with any
interpretation as yet offered except that which refers the change
to the accumulation by selection of variations which promote a
likeness to the Danaidas.
The facts also bear upon the two theories of Mimicry associated
with the names of H. W. Bates and Fritz Miller. According to
Bates’ theory, Mimicry is a special form of protective or cryptic
resemblance. In the ordinary examples of this principle, species
are aided in the struggle by concealment, by a likeness to some
objects of no interest to their enemies (such as bark, earth, etc.) ;
in these special examples (called mimetic) species are aided by
resembling some object which is unpleasant or even dangerous
to their foes. Fritz Muller’s theory of Mimicry includes the
cases in which the mimics, as well as their models, are specially
defended, although generally to an unequal degree. The”
resemblance is due to the advantages of a common advertise-
21 In a letter to Sir Charles Lyell, Oct. 11, 1859.—Lzfe and Letters, ii. 212.
* It is probable that relative abundance may determine the relationship of
model and mimic in cases where there is no reason for suspecting any difference in
the degree of unpalatability.
No
-
1909] Mimicry in the Butterflies of North America 2
ment. Before the growth of a mimetic likeness, Batesian
mimics, it is reasonable to assume, belonged to the immense
group of species possessing a cryptic appearance; Mullerian
mimics on the other hand may be assumed to have possessed
warning or aposematic colours of their own previous to the
adoption of those of another species. This test is more readily
apphed than might be supposed; for a comparison with allied
non-mimetic species, and with the non-mimetic males of mimetic
females, will generally indicate whether the ancestral pattern of
a species now mimetic belonged to the group of concealing colours
or to that of warning.
The Danaidas invaded North America and entered an assem-
blage of butterflies of which the dominant species are ancient
inhabitants of the northern land-belt. Among them are sev-
eral such as the species of Grapta or Polygonia (the ‘‘Comma”’
butterflies), with beautifully cryptic patterns on the parts of the
wing surface exposed in the resting position. No such forms
have been influenced by the invaders, but with the whole fauna
before them they have only produced changes in the dominant
group Limenitis, known throughout the northern belt for a con-
spicuous under surface and a floating flight; also believed to be
mimicked by other butterflies, e. g., the females of the Apaturas
(‘Purple Emperors’’) and the later brood of Araschnia levana.”
Furthermore, the close allies of Limenitis in South America, the
abundant Adelphas, are beautifully mimicked, not only by
females of the genus Chlorippe, which represents A patura, but
also by Erycimdae. In another point the facts are at variance
with Bates’ interpretation but harmonize with Muller’s. Bates
supposed Mimicry to be an adaptation by which a scarce, hard-
pressed form is enabled to hold its own in the struggle for exis-
tence But L. arthemis, which represents with little or no change
the species from which the mimics were derived, persists as a very
abundant and flourishing species, while its mimetic descendant
archippus has gained an immensely extended range and become
almost universally commoner than any other species of its group
(Scudder, 1. c. 266). L. archippus extends from Hudson’s Bay
to the Gulf of Mexico; over this vast ared it is only rare in the
west, and only unknown in Colorado, Arizona, and New Mexico
(l. c. 278). It is to be observed that the range of archippus in-
cludes the whole of the area (Canada and the north-eastern States)
occupied by the ancestral form arthemis.
3 See also the mimetic resemblance to L. astyanax described on pp. 229-30.
Annals Entomological Society of America [Vol. II,
bo
bo
iS)
The facts indicate that the changes produced by the invaders
were wrought in the conspicuous pattern of a dominant indi-
genous species, and that the transformed butterfly having adopted
the advertisement of the still more unpalatable Danaida, became
even more dominant and gained a far wider range than before.
The mimetic resemblance arose in a species which we have reason
to believe possessed warning colours and some form of special
protection before the change occurred. There is no evidence that
the special protection was diminished after the assumption of
Mimicry, and, if it remain, the new appearance is still a warning
character, only one that is learnt by enemies more readily than
the old because of the wide advertisement given to it by Danaida
plexippus. The facts harmonize with the theory of Fritz Muller
rather than with that of H. W. Bates.
THE ‘PoIsoN-EATING’ SWALLOW-TAIL BUTTERFLIES (PHAR-
MACOPHAGUS) AS MODELS FOR MIMICRY.
The late Erich Haase gave the name of Pharmacophagus or
‘‘Poison-eater’’ to the section of swallow-tail butterflies whose
larvae feed upon Artstolochia or allied species, and he made the
probable suggestion that the qualities which render them dis-
tasteful are derived from the juices of the food-plant. The poti-
son-eating swallow-tails are abundant in tropical America and
the Oriental Region, but with the exception of antenor in Mada-
gascar are wanting from the Ethiopian Region. They are exten-
sively mimicked by swallow-tails of the other two sections:
Papilio, of which machaon may be taken as a type, and Cosmo-
desmus, of which podalirius serves as an example. The distinc-
tion between these three sections of Papilionidae extends to lar-
val and pupal stages, as was originally discovered by Horsfield.
It was made the basis of Haase’s classification,”* recently confirmed
and amplified by Rothschild and Jordan.” The latter authorities
propose the names ‘‘Aristolochia Swallow-tails,” ‘‘Fluted Swal-
low-tails,”’ and “‘Kite Swallow-tails,’’ respectively for Haase’s
sections Pharmacophagus, Papilio and Cosmodesmus.
The Pharmacophagus swallow-tails are not so well-known as
models for Mimicry as are the Danatinae, Acraeinae, etc., and it is
therefore expedient to say a few words about the section before
considering the effect produced by one of its members in North
America.
24 Researches on Mimuicry, Pt. ii, Stuttgart, 1896, English translation.
25 Nov. Zool. xiii (1906), 411-752.
to
1909 | Mimicry in the Butterflies of North America 23
In tropical America not only are the species of Pharmaco-
phagus extensively mimicked but Mimicry is also strongly devel-
oped within the limits of the section itself, viz., between the
two dominant groups Aeneas and Lysander. In these groups
the males are commonly very different in appearance from the
females and frequent more open habitats such as the banks of
rivers, etc., the females being found in the forest. In the inter-
nal Mimicry between Aeneas and Lysander the males resemble
the males, the females the females, but the female patterns are
alone extensively mimicked by other groups—Papilio, Cosmo-
desmus and certain Pierinae. I have as yet only come across a
single example (a Cosmodesmus) in which the pattern and green
markings of the males are mimicked. One or two species (e. g.
Ph. hahnelt) of Pharmacoghagus are themselves mimics of domi-
nant Ithomiine genera.
It is well known that in the Papilio mimics of Pharma-
cophagus the resemblance is often attained by the females alone,
a tendency exemplified in North America as shown on pp. 224-6.
In Cosmodesmus, on the other hand, where the Mimicry of these
models reaches a far higher level of perfection, it is equally pro-
nounced in both sexes. In Africa, on the other hand, where,
in default of Pharmacophagus models, the swallow-tails of both
groups frequently mimic Danainae and Acraeinae, the resem-
blances obtained by Cosmodesmus are far less striking than those
of the other section; yet the relationship of Mimicry to sex re-
mains unchanged.
In the Oriental Region the female Mimicry of Pharmacophagus
is still characteristic of Papilio, also appearing in certain Cosmo-
desmus mimics of Danainae. Two remarkable features appear
in this Region: (1) the development within Pharmacophagus
of the gigantic Ornithopteras which do not appear to be mim-
icked at all; (2) the appearance within the section Papilio of
groups which are mimicked as extensively, perhaps even more
extensively, than Pharmacophagus itself. Among the mimics of
these Papilios are not only species of other proups in the same
section but also, although in small proportion, Satyrine butter-
flies and day-flying moths.
The fact that Pharmacophagus and certain groups of Papilio
should be mimicked pre-eminently by other Papilionidae is evi-
dence that Mimicry is most easily attained when there are initial
resemblances of size, shape, habits, and modes of flight upon
which to build. as 4
to
24 Annals Entomological Society of America [Vets Tt,
PHARMACOPHAGUS (PAPILIO) PHILENOR L., AS A MODEL FOR
Mrimicry IN NortH AMERICA.
Pharmacophagus is a tropical assemblage, but a few species
have found their way into the northern belt in both the Old
World and the New. Pharm. polydamas, with an immense range
in South and Central America, also extends into the northern con-
tinent but does not there become the object of Mimicry. Pharm.
philenor, ranging through Mexico and the United States (except
the central district from Colorado northwards) but only as a
straggler in New England and southern Canada, is on the other
hand an important model for Mimicry.
There is here no such interesting history of past migrations
to unfold as we were able to trace in the American Danaidas.
Ph. philenor is a member of the distinctively New World species
of Pharmacophagus associated together and separated from the
Old World species by structural characters. Rothschild and
Jordan state that every species can be recognized as American
by the examination of a single joint of one leg, and they are
therefore justified in concluding that all the New World species
were derived from a single ancestor possessing this character.
There is no sufficient evidence that any of the numerous patterns
are ancestral as compared with the others, although it is tolerably
safe to conclude that the presence of hind-wing ‘‘tails”’ 1s primi-
tive as compared with their absence. Following this indication,
we find that as a general rule the specialized and modern forms are
predominant nearer to the Equator, the comparatively ancestral
tailed forms occurring in latitudes more remote from it both north
and south.
Ph. philenor is a ‘‘tailed”’ form, although its sub-species orsua
in the Tres Marias Islands is nearly tailless. It is probably an
intruder into North America from the tropics of the same Contt-
nent. It is well known to possess the characteristics of distaste-
ful species—gregarious larvae, tenacity of life, and a strong, dis-
agreeable scent.
oe
Tue THREE Papitio Mimics oF PH. PHILENOR IN NORTH
AMERICA.
The three swallow-tail mimics of philenor belong to separate
groups of Haase’s section Papilio. All of them range from the
Atlantic to the Mississippi basin.
190g] Mimicry in the Butterflies of North America 225
The female of Papilio polyxenes astertus (Cr.) belonging to the
MacuHaon Group mimics philenor on both surfaces, the male on
the under surface alone, except at Guerrero, Mexico, where a form
(ampliata) mimetic on the upper surface is transitional into the
ordinary male.
Papilio glaucus glaucus (L.) belongs to the GLaucus Group,
next but one to the group containing asterius.** The female is
dimorphic, one form resembling the male and the other, the turnus
form, mimetic of philenor (L.), becoming commoner in the south-
ern part of the range. In the closely allied sub-species P. glau-
cus canadensis (Rothsch. and Jord.), the mimetic female form is
unknown.
Paptlio troilus troilus (L.) belongs to the next succeeding
Troi_us Group, allied to the tropical and highly mimetic ANn-
CHISIADES GROuP, with gregarious larvae. Both male and female
of trotlus mimic philenor on both wing surfaces.
The most remarkable fact about these three mimics is not
their moderate resemblance to the primary model philenor, but
their extraordinary likeness to one another. Upon the wing or
at rest at a little distance they would be indistinguishable, -and
even in the cabinet they may be easily confused. It is to be
expected that the species of allied groups, with patterns converg-
ing towards that of a single model, and approaching it by varia-
tions which tend to be produced in the section to which they
belong, should incidentally approach one another. But the
strong likeness between the mimetic forms of trodlus asterius, and
glaucus seems to require something more than this, and supports
the conclusion that there is secondary Mimicry between the mim-
ics themselves. It is not necessary to repeat here the details of
these secondary resemblances,?’? and as a matter of fact the like-
ness itself 1s stronger than might be inferred from a consideration
of the details themselves. It is necessary to see it in order to
appreciate it.
It is probable that trozlus, mimetic in both sexes, is the oldest
mimic; astertus, non-mimetic on the upper surface of the male or
with very rough incipient Mimicry, the next to appear; and glau-
cus, mimetic in only one form of the female, the youngest. These
conclusions as to relative age are on the whole supported by the
relative strength of the detailed resemblances to philenor in the
three mimics.
°° The species is commonly called P. turnus and its mimetic female the
glaucus form. I follow Rothschild and Jordan in transposing these names.
27 See Trans. Ent. Soc. Lond. (1908), 467-471.
Annals Entomological Society of America [Vol mule
to
to
Ov
In attempting to trace the past history, here again we have
the great advantage of knowing the more ancestral patterns
from which the three mimics were derived :—troilus from a pala-
medes-like form; asterius from the pattern of its male, which again
leads back to the typical pattern of the MacHAaon Group: the
turnus female of glaucus from the male and non-mimetic female
of the same species.
It is highly probable that the earliest steps in the direction of
Mimicry in astertus and glaucus were favoured by the appearance
of partially melanic varieties of the female, thus effecting sudden-
ly that essential change which enables a butterfly with a yellow
ground-colour to become the mimic of one in which it is black.
But this transformation, immensely important as it is, supplies
nothing more than a tinted paper for a new picture. That the
melanic varieties were partial is clearly shown by the persistence
(in glaucus) in a subdued and inconspicuous form of certain
ancestral features that do not contribute to the Mimicry, but
above all by the retention of every element in the original pattern
that can be worked up into the new. By the modification of these
elements in form or colour,—often in both form and colour,—the
detailed mimetic pattern has been wrought upon the darkened
surface.
Valuable confirmation of the history suggested in the last
paragraph is to be found in the dark form melasina (Rothsch.
and Jord.) found in both sexes of P. polyxenes americus (Kollar),
extending from North Peru to Colombia and Venezuela. This
melanic variety probably represents the darkened form of asterius
before the initiation of the detailed mimicry of philenor. The
sub-species americus does not enter the range of philenor, and
those ancestral elements which have been retained by its melanic
form have not developed into the mimetic likeness seen in the
more northern sub-species asterius.
It is well known that all four species (including philenor) fly
together. Even in my own limited experience I have taken three
of them in adjacent streets on the outskirts of Chicago on the
same day (Aug. 10, 1897), and the fourth in the same locality a
little earlier (July 28). But precise knowledge of their relative
proportions in different parts of their range would be of highest
interest. Again, troiulus extends to the North-West Territory
of Canada, probably far beyond the area in which philenor occurs
as a straggler; and it would be very interesting to compare min-
1909] Mimicry in the Butterflies of North America 227
utely large numbers of such specimens with those from districts
where the model is dominant. <A similar study should be made
of the Canadian specimens of asterius, although this species does
not extend so far beyond the northern limits of the poison-eating
model.
From another point of view the interbreeding of the turnus
female of glaucus with a male from some northern district where
turnus is unknown or very scarce would be of the highest interest.
We should here be able to test whether the Mendelian relationship
exists between the parent form and its partially melanic variety
further transformed by selection,—not a mere melanic ‘‘muta-
tion.’ trust that my friend Prof. CB. Davenport may be
able to undertake this experiment at the Cold Spring Experi-
mental Station. I cannot doubt that breeding could be easily
carried through two generations in a large enclosed space exposed
to the sun and planted with abundant flowers and the food-plant
of the species. It would probably be safe to use Long Island
males, while female pupae or the freshly bred females themselves
could be readily obtained from further south.
THE EvoLuTion or LIMENITIS (B.) ASTYANAX (F). as A Mimic
oF PH. PHILENOR AND Its Papitio MIMICcs.
Scudder states that L. astyanax “‘ranges from the Atlantic
westward to the Mississippi Valley, and from the Gulf of Mexico
northward to about the 43rd parallel of latitude.’’’* It thus falls
entirely within the area of philenor. The northern boundary
of astyanax corresponds with the southern limit of its parent
arthemis, and Scudder (1. c. 289) considers that they interbreed
and that the intermediate form proserpina, found along the nar-
row belt where the two species or sub-species meet, is the result-
ing hybrid. Both arthemis and proserpina have been bred from
the eggs of the latter. There seems lhttle doubt that astyanax 1s
a very recent development from arthemis in the southern part of
its range,—so recent that the areas of distribution still remain
distinct and parent and offspring only meet along a narrow line.
It is probable that archippus arose in the same manner in part
of the area of arthemtis, but that later, after the separation had
become complete, it spread northward over the whole range of
its parent.
28 A closely allied species or probably a form of the same species is recorded
by Godman and Salvin from Mexico.
228 Annals Entomological Society of America [Vol. II,
The evolution of astyanax from arthemts was far simpler than
that of archippus. The great difference in appearance between
parent and offspring is brought about, as regards the upper sur-
face, by the disappearance of the broad white band of arthemis
together with all but a trace of the sub-apical white markings of
the fore wings. Over and within the area formerly occupied by
the white band, a bluish or greenish iridescence spreads from the
marginal region where it exists in arthemis. This marginal irides-
cence—Just as in astyanax—is bluish in some individuals of arthe-
mis, greenish in others. Reddish sub-marginal spots, although
rarer in the hind wing of astyanax, are actually commoner in
the fore wings than in arthemis. This curious fact, together
with the evidence that astyanax and archippus may occasionally
interbreed, suggests the possibility of some connection between
the origins of the two mimics.
The under surface of astyanax has not only similarly lost the
white markings, but the chocolate-brown ground-colour of arthe-
mis has become transformed into a ‘ark iridescent greenish-
brown. Against this background the reddish spots near the
margin and base of the wings become far more conspicuous than
in the parent form. The material for this transformation in tint
is still to be seen in the great variation of the ground-colour in
arthemts.
Although, as Scudder rightly maintains (1. c. 287), L. astyanax
is a very poor mimic of Pharm. plilenor, it bears considerable
resemblance to the three Papilio mimics, especially trodlus.
Although the iridescent blue or green of its upper surface ap-
proaches rather more closely than the Papilios to the brilliant,
steely lustre of philenor, it is still in this respect widely separated
from the primary model and near to the mimics. The reddish
spots of the under surface offer but a rough likeness to those of
any of the above named species, but there can be no doubt that
their emphasis is an element in the mimetic resemblance.
A careful examination of large numbers of astyanax from the
extreme south of the range where it passes out of the area of
glaucus and troilus but remains within that of philenor and aster-
dus, might yield interesting results. An investigation of the pro-
portion it bears to the four Papilionidae in various parts of their
common range would also be of deep interest. Of the highest
importance would be the attempt—which would probably be
successful—to breed astyanax and arthemis and to ascertain
1909] Mimicry in the Butterflies of North America 220
whether the Mendelian proportions appear in the offspring of the
hybrids. The pairing of astyanax and archippus, although in
this case failure is probable, ought also to be attempted.
THE FEMALE OF ARGYNNIS (SEMNOPSYCHE) DIANA (CR.) A
Mimic oF LIMENITIS ASTYANAX.
The comparatively narrow range of this species is, as Scudder
points out, wholly included within that of astyanax (1. c. 1802).
The Mimicry is confined to the upper surface, where the blue
tint has even less sheen than that of any other member of the
group clustered round the brilliant philenor. Apart from the
blue expanse, which he admits to be mimetic, Dr. F. A. Dixey
considers that the female of dzana belongs to a set of dark female
forms well-known in Argynnis, forms which he believes to be
ancestral.2® It is probable that ‘the recent evolution of L.
astyanax provided this ancestral form with a model which it could
approach by small and easy steps of variation.’*°
THE BEARING UPON THEORIES OF MIMICRY OF PHARM.
PHILENOR AND Its MuImICcs.
Haase, who always shows an imperfect appreciation of the
scope of Fritz Miller’s principle, apparently regarded all the
species mentioned in the preceding section as simple Batesian
mumics, of philenor, neglecting the mimetic relationships between
the mimics themselves. This interpretation is unconvincing,
and most naturalists will agree with Scudder in his hesitation to
accept the two Nymphalines, astyanax and diana (female),
as simple mimics of philenor. The Mullerian hypothesis at once
explains relationships that are mere coincidences under that of
Bates.
Pharm. philenor, a probable intruder from the American
tropics, produced its effect upon the three large Papilios—butter-
flies with a conspicuous under surface pattern, in large part
reproducing that of the upper surface, butterflies belonging to a
section that provides models for extensive Mimicry in the Oriental
Region. They may be regarded as Mullerian mimics of the pri-
mary Pharmacophagus model, exhibiting a certain amount of
Secondary Mimicry of one another.
The four above-named Papzilionidae, but especially the three
mimics acting as secondary models, then proudced an effect upon
29 Trans. Ent. Soc. Lond. (1890), 89-129.
80 Ibid., (1908), 475.
230 Annals Entomological Society of America [Vol EL,
L. arthemis—that same conspicuous, specially defended element
in the North American butterfly fauna which was influenced in
an entirely different direction by the Danaine invaders. The
result of the former influence is seen in L. astyanax, a secondary
mimic of the three Papilio mimics of philenor.
One of the most interesting elements in this complex mimetic
system is the final appearance of a tertiary mimic of astyanax,
viz., the female of Argynnis diana. This was recognized by
Scudder, although, not fully appreciating the Mullerian hypo-
thesis, he was much puzzled by the fact.*
The under surface of the female dzana is inconspicuous, and,
considering also the restricted range and relative rarity of the
species, 1t is probable that this member of the assemblage of
species convergent round plilenor isa Batesian mimic. But its
resemblance to astyanax supports the conclusion that this latter
and the sister-species archippus (and its forms) are Mullerian
mimics and the parent arthemis a specially protected species.
The resemblance of astyanax to the three species of the section
Papilio, as well as the secondary resemblances between the three,
similarly supports the conclusion that these mimics are Mullerian.
I have not hitherto called attention to the paramount need for
experimental research and field observations directed to test for
the presence of distasteful qualities and to estimate their effect
upon enemies of the most varied kinds. It 1s of the utmost
importance that such investigations should be undertaken on the
largest possible scale. In the meantime the Mullerian Hypothe-
Sis appears to explain a series of remarkable relationships which
remain coincidences under any other hypothesis.
THE RESEMBLANCES BETWEEN LIMENITIS (ADELPHA) CALI-
FORNICA (BuTL.) and LimENITIs (NAJAS) LORQUINI
(Boisp.)
The examples of Mimicry which we have been considering
hitherto are, with the exception of the widespread L. archippus,
characteristic of the eastern side of North America. The present
instance, the last of the examples known in this portion of the
northern land-belt, is found on the Pacific coast. The resem-
blances are somewhat crude but of quite remarkable interest.
Limemitis californica, because of its pattern and colouring, is
often placed in Adelpha, a large genus with over seventy species
31]. c. 718, 1802; see, however, 266, where Scudder suggests that astyanax
may possibly be specially protected.
1909 | Mimicry in the Butterflies of North America 231
all confined to tropical America. Adelpha is separated from the
closely allied northern genus Limenitis by the hairiness of the
eyes in front. Caltfornica is by this character as well as its more
northern range associated with the heterogeneous assemblage
‘‘Limenitts,”” which so much requires a thorough revision. In
adopting this view I accept the position assigned to the species
by ocudder im 137 5.°7
Closely allied to californica, of Oregon, California, and Nevada,
is L. bredowt (Hutbn.) of Arizona, Mexico, and Guatemala. A
much needed investigation is the determination whether these
two forms meet, and interbreed along the line of contact.
The southern species or sub-species bredowz, 1s associated in
Mexico and Guatemala with many true species of Adelpha of
which no less than thirty-one extend into Central America. To
these it, and to a less extent the northern californica, bear much
likeness, especially to A. dyonysa (Hew.), masstlia (Feld), lerna
(Hew.), and fessonia (Hew.). This likeness is probably a mime-
tic resemblance which extends beyond the range of the models
into Arizona, and, with diminished effect, still further north into
the allied sub-species. _Although the details of the resemblance
leave little doubt that this interpretation is correct for the south-
ern bredowt, it 1s possible that californica represents an ancestral
form connecting the Adelphas with Limenitis, a form left isolated
and comparatively unchanged in the north,* while its southern
allies have been modified by the presence of the dominant
Adelphas. At any rate in one feature neither sub-species appear
to be mimetic, viz., in the yellowish tint of the conspicuous band
crossing both wings; for in all the Central American Adelphas
at all resembling them, this marking is pure white or bluish-white.
We cannot hope to determine how far the pattern of californica
is ancestral until the structural relationships and the early stages
of Limenitis in the widest sense and Adelpha have been ‘most
minutely investigated. .
Limemitis lorquint, occurring with L. californica in Nevada,
California, and Oregon, also extends far north of this species into
British Columbia and Vancouver Island. Among all the North
American species of Limenitis it is the one which comes nearest
to the Old World forms, as Scudder recognized when he included
it with the European L. populi in the genus Najas, separating all
3 Bull. Buffalo Soc. N. Sc. (Feb., 1875), 233.
33 See, however, p. 234-5.
222 Annals Entomological Society of America [Vol. II,
the other American forms of Limenitis except californica as Basil-
archia. Even such fleeting characters as the markings show the
Old World affinities of lorquini in the strong development of the
pale spot in the fore wing cell and the position and form of the
pale band crossing both wings. It is to be noted furthermore
that its distribution, and especially its extension northward,
along the Pacific coast, bring lorquint into closest proximity to
the Old World species.
In certain important respects the upper surface pattern of
L. lorquint is certainly mimetic of californica: —
The conspicuous fulvous apical area of the fore wing; the yel-
lowish tint of the band crossing both wings, and, although here
the interpretation is less certain, the fulvous marking at the anal
angle of the hind wing.
r. In the first and most important of these points of super-
ficial resemblance there is, so far as my experience goes, a much
greater average development of the fulvous patch in specimens
of lorquint which enter the range of californica in Oregon and
California than in those which come from Canada, entirely beyond
the range of the model.
The close relationship between californica and lorquint may
incline naturalists to look on their resemblance as due to affinity
and not to Mimicry. ‘“‘It is commonly forgotten that Mimicry,
being zndependent of affinity, occurs between forms of all degrees
of relationship, the closest as well as the most remote;’’*4 although
of course the latter are easy to interpret, while the former may be
excessively difficult. In this case, however, there is neither doubt
nor difficulty, for not only is there the geographical coincidence
between the model and the average increase of the marking in the
mimic, but the fulvous apical marking of lorquini—of a somewhat
richer, deeper shade than the tawny patch of californica—is due
to the inward growth of a marginal marking, while that of the
model occupies a clearly defined sub-marginal and sub-apical
position. The resemblance is, in fact, produced by markings
which are essentially different; yet in some of the southern exam-
ples of lorquint in which the markings extend inward to the
greatest distance the superficial resemblance is very considerable.
The above stated conclusion that the chief mimetic element
of lorquint is on the average subject to considerable strengthen-
ing in the southern part of its range, is founded on an examination
34 Trans. Ent. Soc. Lond. (1908), 482.
Ww
W
1909 | Mimicry in the Butterflies of North America 2:
of the few dozen specimens I have been able to study in English
collections, and especially the Godman-Salvin material in the
British Museum. I now trust that the subject may be taken up
by American naturalists and many hundreds of specimens com-
pared from all parts of the north and south range of the species.
2. Inthe second point also, the yellowish tint of the principal
band, the resemblance is certainly mimetic and not due to affin-
ity; for lorquint, ancestral in certain other features, has here lost
the original whiteness of this marking, preserved not only in the
Old World but in Limenitis arthemis and L. weidermeyert (Edw.)
of the New. An excessively slight deepening of the yellow tint
could be made out in southern individuals from the area occupied
by the model. In order to detect the difference, a long series
of northern specimens should be placed beside a similar series
from the south and the two compared in a strong light. But
far larger numbers than I have seen ought to be examined from
this point of view, and, if it were possible to make it, the compari-
son of perfectly fresh specimens would be most desirable.
3. The fulvous marking at the anal angle of the hind wing
is excessively variable and often absent from specimens in all
parts of the range. The comparison of a very large amount of
material is necessary before we can reach any safe conclusions
as to the existence of mimetic resemblance in this feature, and the
same is true of the extremely variable under surface of lorquint,
in which the development of the inner row of sub-marginal bluish
lunules may be mimetic of californica. This feature was gener-
ally suppressed in the Vancouver Island specimens I have seen.
We now come to the consideration of certain differences
between L. californica and its southern form bredow1 which pro-
mote a likeness to lorquini. If these are not mere coincidences,
we can hardly escape the conclusion that there is Reciprocal
Mimicry (Diaposematism) between californica and bredow.
tr. The wings of both sexes of californica are more rounded
than those of the males of bredow, in this respect resembling both |
sexes of lorquint. The fact that the southern females have
rounded wings may indicate that this character is ancestral in
both sexes, the males alone having been modified in Mimicry of
Adelpha. But it is a probable hypothesis that the presence of
lorquint has prevented this mimetic feature from passing north-
ward into the males of californica. It does pass far beyond
Adelpha in the northernmost part of the range of bredowz in
Arizona.
234 Annals Entomological Society of America [Viol iT;
2. The fulvous marking at the anal angle of the hind wing
which forms so characteristic a feature of bredowz, is greatly re-
duced in californica, approximating to lorquini, which in this
respect may be advancing to meet its model (see p. 233.)
3. The following points concern the band crossing the fore-
wing. Owing to the small size of the last spot in caltfornica
and the different direction of the spot next to it, the junction
between the bands of fore and hind wing forms a step-like break
in californica, whereas 1n bredowr the bands tend to be continuous,
approximating more closely to the single smooth streak crossing
both wings in the Adelphas. In lorquinz this step-like break and
want of continuity in direction is even more pronounced. Again,
the fore wing band of lorquini—one of its ancestral features—
forms with the adjacent hind wing spot, a drawn-out zigzag like
a flattened-down W. By a modification in the position and
direction of the spots of californica as compared with bredow,
it also gains the appearance of a very flattened W, although a far
less regular one than that of lorquimi. The resemblance is
only superficial; for corresponding spots do not occupy the
upper angle of the W in the two species. But the attainment of a
likeness by means that are different from those employed in
another species supports the interpretation of the resemblance
as mimetic.
Whatever be the true interpretation of the resemblances above
described, it is of the utmost importance and interest to study
the relative numbers of calzfornica and lorquint at as many differ-
ent points as possible in their common range, to observe how far
they fly together and present the same appearance on the wing
and at rest from a little distance, and to test their relative pala-
tability on a variety of insect-eating animals found in the same
area.
The following general considerations support the conclusion
that californica is not an ancient element in the Pacific fauna of
North America, but a comparatively recent intruder from the
south—an intruder that has modified the indigenous inhabitant
lorquint and has been also reciprocally modified thereby.
Limenitis in the broad sense is part of the ancient northern
butterfly fauna of North America. It has here split up into sev-
eral well-marked species characteristic of the area. It is highly
susceptible to mimetic influence—far more so than any other
North American group—and contributes the majority of the
iS)
ios)
wat
T909| Mimuicry in the ‘Butterflies of North America
examples of Mimicry from this part of the world. L. archippus
has been shown to be the result of a recent invasion,—its southern
and eastern forms to be still newer products of the changes in
archippus itself. .
The sensitiveness of the group is shown by the fact that, in
spite of this recent origin, all except astyanax are most beautiful
and striking mimics; and even astyanax is a better mimic than
lorquint. The fact that lorquint, the member of so sensitive a
group, is an undoubted mimic, but a very poor mimic, supports
the conclusion that the association with its model has endured for
but a brief period, a conclusion also supported by the diminution
of the resemblance outside the range of californica.
If the relationships which I have found to exist in the avail-
able material—in quantity very insufficient for such minute com-
parisons—if these are confirmed by extensive investigations in
America, it will follow that the resemblances between L. calzfor-
nica and L. lorquini will be one of the most interesting and instruc-
tive examples of Mimicry in the world. Its value will lie in the
early stage reached by the resemblance, together with the
diminution of the likeness in californica to the south and, especial-
ly, in lorquint to the north. There is no reasonable doubt that
lorquint forms a single Syngamic community along the Pacific
Coast of North America, and we should therefore witness, first,
the marked strengthening of characters in an area of selection;
secondly, their transmission with diminished effect into other
areas.
If what I have observed be the phenomena presented by the
growth, at an early stage, of a mimetic likeness in lorquini, then
that growth is ‘‘continuous” and transitional to the last and
finest decree:
It is perhaps appropriate to’ state in a few lines how we may
imagine that the selection of minute characteristics such as the
presence or the position of a single spot may be made. We
ourselves may observe that one individual butterfly is a better
mimic than another. We may then analyse the pattern, as I
have attempted to do in this address, and realize that the
improvement is due to differences in one or more relatively minute
elements. Recognizing the cause of the change, we are perhaps
prone erroneously to suppose that enemies recognize it also and
that selection has been brought to bear directly and consciously
upon it. Such a view is almost certainly wrong. The only pro-
236 Annals Entomological Society of America [Vol. II,
bable hypothesis is that sharpsighted enemies, without analysing
the markings, recognize differences in degrees of likeness, and
that the selective pressure exercised by them is influenced by
the recognition.
A great deal of attention is rightly directed at the present
day to the value of experiment, and indeed it is impossible to
over-estimate its importance. But while human performance is
of the deepest interest for the solution of mysteries innumerable,
of more profound significance still, for the comprehension of the
method of evolution, is the vast performance of Nature herself.»
Because of the bright promise it holds for the understanding of
Nature’s experiments, I have brought before you the subject of
Mimicry in North American Butterflies.
In the introductory words I spoke of the relationship of my
subject to the teachings of Darwin, and now I am anxious to con-
nect this address by a closer link to the personality of the illus-
trious naturalist. With the kind consent of Mr. Francis Darwin,
I am able to achieve this object by printing, for the first time, a
letter, recently discovered in the archives of the Hope Depart-
ment of Oxford, written by Darwin to the Founder in 1837. It
is concerned with the insect material collected on the Beagle
and is of peculiar interest because so few of Darwin’s letters of
this early date have been preserved. The letter clearly exhibits
the keen interest which Darwin took in the working out of his
collections, and the free and generous use he made of his material.
A number of Diptera captured by him in Australia and Tasmania
—evidently gifts to Mr. Hope—exist in the Hope Department,
and are still in excellent condition. It is probable that species
of other groups collected by .him are also present.
DEAR HOPE
I called yesterday on you and left a tin box with a few Hobart
Town beetles, which I had neglected to put with the others. Is not
there not [sic] a Chrysomela among them, very like the English species
which feeds on the Broom.—I have spoken to Waterhouse about the
Australian insects; you can have them when you like.—The collections
in the pill boxes come from Sydney, Hobart town, and King George’s
Sound.—Do you want all orders for your work? Some are already I
believe in the hands of Mr. Walker, and you know Waterhouse has
described some minute Coleoptera in the papers read to the Entomo-
35 See Carl H. Eigenmann in Fifty Years of Darwinism, New York (1909),
208.
1909] Mimacry in the Butterflies of North America 234
logical Soc: To these descriptions of course you will refer —You will
be glad to find that many of the minute Coleoptera from Sydney are
mounted on cards.—Will you send me as soon as you conveniently
can, one of my boxes, as I am in want of them to transplant some
more insects.—Perhaps you had better return the Carabi, as they came
from several localities I am afraid of some mistake. We must put out
specimens for the Entomolog: Soc.: and your Cabinet. May I state
in a note on your authority that a third or a half of the insects which
you already have of mine from Sydney and Hobart town are undes-
cribed.—It is a striking fact, if such is the case, for it shows how imper-
fectly known the insects are, even in the close neighborhood of the
two Australian Capitals.
Floreat Entomologia
Yours most truly,
Wednesday. Cuas. DARWIN.*
The last words of Darwin’s letter are surely a most fitting
conclusion to this Anniversary Address, and I conclude by quoting
his humorous repetition of them probably twenty years later.
“ “Floreat Entomologia’!—to which toast at Cambridge I have
drunk many a glass of wine. So again, ‘Floreat Entomologia.’
N.B. I have not now been drinking any glasses full of wine.*7
CONCLUSIONS.
It will probably be convenient to sum up rather fully the chief
conclusions contained in the foregoing address.
1. The study of Mimicry possesses special advantages for
an understanding of the history and causes of evolution.
2. North America is the most suitable area in the world in
which to begin the study of Mimicry.
3. The great American Danaine butterflies, formerly includ-
ed in the genera Anosta and Tasitia, are a foreign element in the
New World fauna. They bear the closest affinity to a large group
of indigenous Old World Danainae, and should be fused with the
nearest of these (Limnas, and Salatura) into a single genus,
Danaida.
* The letter is addressed: ‘‘The Revd. F. W. Hope, 56, Upper Seymour
Street.” At the head Mr. Hope had written ‘‘D,” and the date ‘'1837.”’ The
red-stamped post-mark gives the date ‘‘Ju. 22, 1837.’’ Darwin’s own address
(36, Great Marlborough Street) does not appear. At the date of the letter the
Entomological Society of London possessed a large collection of insects, long
since dispersed. Darwin knew Mr. Hope before the Voyage, and speaks in let-
ters to W. D. Fox (1829-30) of his splendid collection and of his generosity with
specimens. He also went for an entomological trip in North Wales with Hope
(June, 1829), unfortunately broken short for Darwin by ill health. See Life
and Lettérs, i. 174, 175, 178, 181. G.°R. Waterhouse and Francis Walker,
reterred to in the letter, were both on the staff of the British Museum.
To Sir John Lubbock (Lord Avebury), some date before 1857.—Life and
Letters ii. 141.
238 Annals Entomological Society of America [ Vol-atle
4. The Old World origin of Danaida is also proved by the
extent and variety of its mimetic relationships; while the path
of its invasion of the New World and of South by way of North
America, may be traced by foot-prints, as it were, of mimetic
erect.
5. That Danaida plexippus is the older invader is equally
shown by the depth of the impression it has made and the
amount of change it has itself undergone in the New World.
6. Danaida berenice and its form strigosa show comparatively
slight changes in the New World, and, as regards mimetic influ-
ence, have but deepened the foot-prints left by plextppus.
7. Limenitis arthemis, the indigenous ancestor of the mimic
of plexippus, persits with little or no change; and it is possible
to show how far the very different markings of the mimetic
daughter species, L. archippus, have been carved out of those
of the parent.
8. The recent date of this great superficial transformation
is proved by the close resemblances between the larval and pupal
stages of parent and offspring. L. archtppus also probably occa-
sionally interbreeds with the mimetic L. astyanax—a still younger
descendant of the same parent.
9. L. archippus probably arose on the southern borders of
arthenus, but afterwards ranged northwards over the area of the
parent species.
to. The southern astyanax, meeting the northern arthenus
along a narrow belt, is probably repeating the earlier history of
archippus.
11. The forms of sub-species of archippus—floridensis in
Florida and hulsti in Arizona—have arisen from the earlier mimic
of D. plexippus as a result of the predominance in these localities
respectively, of Danaida berenice and its form strigosa.
12. Details of the older Mimicry persist in floridensis (and
perhaps in hulsti), somewhat detracting from the newer resem-
blance.
13. Certain features in the mimetic likeness newly attained
in Florida and Arizona are probably due to the recall or the re-
emphasis of elements in the pattern of arthemts which had been
greatly reduced in archippus.
14. The fact that the invading Danaidas have only influenced
among the whole indigenous butterfly fauna, the dominant con-
spicuous Nymphaline genus Limenitis, supports a Mullerian as
opposed to a Batesian interpretation of the phenomena.
1909 | Mimacry in the Butterflies of North America 230
15. Lhe fact that the ancestral pattern of a species indige-
nous in the temperate zone of the New World should be wholly
transformed by a recent invader from the Old World tropics—
the invader meanwhile retaining its original characteristic pat-
tern, is demonstrative of the inadequacy of the theory which
refers these likenesses to the influence of soil, climate, etc.
16. The poison-eating ‘‘Aristolochia swallow-tail’’ Phar-
macophagus (Papilio) philenor belongs structurally to the Ameri-
can division of this tropical section, and is probably an intruder
into North America from the south.
17. Just as tropical species of Pharmacophagus are mimicked,
especially by other sections of swallow-tails, so the invading
philenor is mimicked by three species of the section ‘‘ Papilio.”
18. Of these three—Papilio troilus, mimetic in both sexes,
is probably the oldest; P. aster1us, mimetic in female and on under
surface of male, the next; and P. glaucus, mimetic in one out of
the two forms of female (the mimetic form becoming more num-
erous in the south of the range), the youngest.
1g. The ancestors of these mimics persist with little or no
change—in the two last-named species, the non-mimetic sex or
form, in the first-named the allied palamedes. By their aid we
can reconstruct the history of the transformation.
20. In astertus and glaucus partially melanic forms of the
female probably supplied a tinted background on which the new
and mimetic picture was gradually built up by the modification
of elements in the original non-mimetic pattern.
21. The close resemblance between the three mimicking
species cannot be entirely explained by their convergence upon a
single model, but seems to imply the existence of Secondary
Mimicry between them.
22. Limemtis astyanax has arisen as a very recent modifica-
tion of arthenus in Mimicry of philenor, and especially in Second-
ary Mimicry of the three Papilio mimics.
23. The female of Argynnis (Semnopsyche) diana has arisen
as a tertiary mimic, on the upper surface, of L. astyanax. Its
under surface, inconspicuous with that of the male when contract-
ed, suggests that the species is palatable as compared with the
rest of this combination and that its Mimicry is Batesian.
24. The dark ground and pale markings of the female diana
are probably analogous with those of other dark female forms
in Argynmdae, while the blue colouring is an additional feature
of purely mimetic significance.
240 Annals Entomological Society of America Voli.
25. The arrangement of the North American butterflies
which converge on Pharm. philenor, in concentric rings each
mimetic of that lying within it, strongly supports a Miullerian
interpretation of all except the species (diana) in the outermost
layer.
26. Laimenitis (Adelpha) californica of the Pacific coast is
probably a Limenitis mimic of the South American genus Adelpha,
to which its southern sub-species bredowz bears a stronger resem-
blance.
27. Limenitis (Najas) lorquint, in some respects the most
ancestral of the North American species of the group, is in other
respects a mimic of L. caltformica.
28. Certain features in which Jorquini superficially resem-
bles californica are on the average more strongly developed in the
area where the two species overlap, while they diminish when
lorquint passes northward of this area.
29. The differences between bredow1, ranging entirely south
of lorquimt, and californica are such as to promote a superficial
resemblance between the latter and lorquim1, supporting the hy-
pothesis that the resemblances between them have been caused
by reciprocal approach (Diaposematism).
30. The differences which distinguish bredowz from californica
are such as to promote a resemblance to the tropical American
genus Adelpha. They are retained by bredowz in Arizona, north
of the range of any true Adelpha.**
31. The detailed study of these resemblances on the Pacific
Coast of North America leads to the conclusion that the Mimicry
is in an incipient stage and that it has been reached and 1s prob-
ably still advancing by minute increments,—that the evolution
is ‘‘continuous”’ to the last degree.
32. In addition to their bearing upon the problems of
Mimicry, the examples considered in the address afford some of
the very best material for testing the operation of Mendel’s Law
under natural conditions.
I wish again to caution my readers that the above conclusions
have been drawn from the careful study of a limited number of
examples. Although insufficient in quantity, the English mater-
ial is as a whole excellent in quality. Thus, many of the Pacific
38 In the southernmost part of the range of bredow?, in Guatemala, the resem-
blance to Adelpha was very slightly augmented in the only two specimens from
this locality I have had the opportunity of studying (Trans. Ent. Soc. Lond.,
1908, 485).
1909 | Mimicry in the Butterflies of North America 241
coast specimens were captured by Lord Walsingham, Dr. Be D:
Godman, and Mr. H. J. Elwes, and the geographical data are of
course as full and precise as we should expect or wish.
I trust that my brother naturalists in America will make a
determined attack on the fascinating problems offered by the
phenomena of Mimicry in the North American butterfly fauna.
In this favoured part of the world the problems have been seen
to be sharp and clear as compared with the almost infinite com-
plexity of the tropics. If my assistance or advice be of any value
it is always at the service of those who desire to undertake such
investigations.
It has been abundantly shown in the course of the address,
that immense numbers of specimens are required from the most
raried localities; and it is likely that difficulties may be presented
by the necessary manipulation, labelling, convenient arrangement
and permanent preservation for the study of future as well as
living naturalists, of so large a mass of material. 1 shall, how-
ever, be most pleased to undertake this part of the investigations
as regards all specimens accompanied by adequate data of space
and time. ‘Such material, preserved in the Hope Department,
may be readily compared with the ever-increasing mass of exam-
ples illustrating the same principles in other parts of the world.
If the indications observed in a small series are still found to
hold in a large one, the growth of such a feature as the orange-
brown apex of the fore wing in Limenttis lorquint would be demon-
strated by a glance at its average condition in specimens from
the different localities as we pass from north to south. Further-
more, we might reasonably hope that a similar series collected
after an interval not greatly prolonged would exhibit differences
in average composition—the actual measurable evidence of the
evolution of a character in a species in the natural state. Even
though such evidence be left for our successors to witness, it still
remains our duty to provide them with the standard by which
alone they will be able to detect and measure it. But Iam hope-
ful of more than this, and think it by no means unlikely that a
part of the reward may be reaped by a single generation of
workers.
An excellent example of work done in a single locality, work
which now requires to be extended to many species in many
localities, is afforded by the data obtained by Mr. J. H. Cook,
and summarized in the following note.
242 Annals Entomological Society of America [Vol. II,
Note.—The capture of males of L. archippus in which the black
stripe was wanting from the upper surface of the hind wing, and of
transitional forms of both sexes at Albany, N. Y., by John H. Cook.
Mr. Cook first met with the stripeless form in June, 1898, near
Hudson, N. Y. A second specimen was captured near his home in
Albany in 1901 and a third 7m the same field in the following year.
This latter was a beautiful specimen apparently only just emerged
from the pupa. Mr. Cook’s attention was now thoroughly aroused
and he collected assiduously at Albany during three seasons, always
working on the best ground to the west of the city, and taking over
90 specimens with the stripe wholly or nearly suppressed. The fol-
lowing conclusions were reached:—(1) All the stripeless archippus
captured were males; (2) The females shared the tendency but never
reached the extreme found in the other sex; (8) Most of the individuals
taken showed some weakening of the stripe, varying from a slight break
(most commonly between veins III and V, and between V, and VII,,
of the system of Comstock and Needham) to complete suppression on
the upper surface; (4) At Albany individuals with a broken stripe
outnumbered those with an entire stripe in the proportion of about
18 to 1, while stripeless specimens were taken in the average propor-
tion of 1 to 14. Mr. Cook also collected data from other localities and
endeavored to interest correspondents in the problem. Including the
Albany material he secured records of about 1600 specimens and was
able to reach the conclusion that in New England and the Middle
States broken-striped individuals are not uncommon though generally
outnumbered by those with a continuous stripe. He did not meet
with any record of a perfectly stripeless form except for his own obser-
vations and the two specimens to which the name pseudodorippus
has been given. Strecker’s type of this form exists in Dr. W. J. Hol-
land’s collection (Butterfly Book, New York [1899], 185). These two
pseudodorippus were also taken in the Eastern States (the Catskill
Mountains, and in Massachusetts), but Mr. Cook, who has seen one and
received from Dr. Holland an account of the other, believes that the
disappearance of the stripe is here part of a general blurring of the
colour-scheme in which some elements are obliterated and there is a
tendency towards the invasion of one colour-area by another. The
extreme varieties captured by Mr. Cook himself, did not, on the other
hand, differ at all from the normal archippus except in the absence of
the black stripe from the upper surface of the hind wings. To this
stripeless variety Mr. Cook and Mr. Watson have given the name
lanthanis. Mr. Cook’s accurate data and most of his specimens were
unfortunately destroyed when the college buildings at Albany were
burnt down on Jan. 6, 1906. It is much to be hoped that he may be
able to continue his most interesting observations in this favourable
locality, and that naturalists may be stimulated, by these reeords,
now by Mr. Cook’s kindness made public for the first time, to work
in other North American localities.
DESCRIPTIONS OF SOME NEW TACHINIDAE.
By CuHaries H. T. TOWNSEND.
The following are descriptions of new forms of Tachinidae
that have been studied during the seasons of 1908 and 1909,
embracing some material handled at the Gipsy Moth Laboratory
in Massachusetts.
Phasmophaga gen. nov.
Front long and very prominent in profile in both sexes.
Front in female about as wide as both eyes, wider in male owing
to the great antennal development in that sex; front and face
nearly equal in width; two to three proclinate orbital bristles
on each side in both sexes, the middle one of the three usually
atrophied. Ocellar bristles strong, outwardly proclinate. Fa-
cialia ciliate half way up, especially ridge-like and pronounced
in male. Frontal bristles descending to insertion of arista in
female, but the arista is inserted higher in the male. No trace
of median carina on facial plate.
Antennae and arista concolorous, wholly orange-yellow; arista
thickened its whole length, a little over half to nearly two-thirds
as long as third antennal joint in male, in female more pointed
and fully as long as third antennal joint, basal joints stout and
as long as wide. Eyes bare. Facial plate very deeply depressed
in male to hold the greatly developed third antennal joint, which
extends almost from plane of front to vibrissae; third antennal
joint in female much more slender and extending less than three-
fourths way to vibrissae. Second antennal joint very short in
both sexes. Vibrissal angles on level with oral margin. Vibris-
sae strong and decussate. Facialia especially bowed in profile
in male, the vibrissae inserted on under side of head. Para-
facials narrower in male than in female. Palpi well developed,
club-shaped. Proboscis short and fleshy. Two sternopleural
and three postsutural bristles. Apical scutellar bristles delicate
but long and decussate. Discal bristles of abdomen present in
both sexes. Hind tibiae with irregular bristles. Apical cell
very short-petiolate or moderately so, ending well before wing
apex. Fourth vein bent at a wide angle, hind cross-vein irregu-
larly bewed or straight, in middle between bend and small cross-
vein.
Reproductive habit, leaf-oviposition. Type, P. antennalis, n.
Sp.
243
.
244 Annals Entomological Society of America [Vol. II,
Phasmophaga antennalis n. sp.
Length, 4.5 to 7 mm. _ Blackish, cinereous pollinose, except the
orange-yellow antennae and arista, the pale yellow palpi, and the red-
dish-yellow legs, of which alone the tarsi are blackish; parafacials more
silvery pollinose. Frontalia nearly equilateral. Arista of male a little
over half as long as third antennal joint. Tegulae whitish, wings clear.
Hind cross-vein irregularly bowed. Apical cell very short-petiolate.
The pollen easily detaches from abdomen leaving it blackish in male,
brownish-yellow on sides and venter in female. From above the last
half of second, third, and fourth abdominal segments appears blackish,
but all pollinose from directly behind.
Type, No. 12618, U. S. N. M.
Two females and three males all reared, with many others,
from Diapheromera femorata, by Messrs. H. H. and H. C. Severin,
the material all being from Wisconsin. The Severins are pre-
paring an exhaustive monograph on the subject of the host and
its parasites.
Phasmophaga meridionalis n. sp.
A collected male, from Plano, Texas, July, 1907 (E. S. Tucker),
may best be placed in this genus, but is a very distinct form from the
preceding species. It has no discal bristles, and the front is short,
not so produced, the frontalia being much widened behind. The arista
is longer, being two-thirds as long as third antennal joint; the petiole
of apical cell is not so short, and hind cross vein is straight. The face
is fully three-fifths of head-width, and the front hardly narrowed.
‘ype, No. 12666, UL 5. N. M:
A male specimen of Phasmophaga in the collection, which
seems same as meridionalis, was reared at Cutler, Florida, from
Antsomorpha buprestoides, May 29, 1908 (Chittenden, No. 361;
Russell, Coll.) It has no discal bristles, the apical cell is short-
petiolate, there are two proclinate orbital bristles, the frontalia
are equal in width and take up fully one-half the width of front.
The arista is scarcely two-thirds the length of the third antennal
joint, the face is about three-fifths of head-width, and the hind
cross-vein is straight.
Dissection of one of the females of P. antennalts, that had been
left in the breeding cage for some days after issuance, disclosed
the characteristic minute eggs of the leaf-ovipositing forms of
Tachinids. It is thus certain that the walking-sticks swallow
the eggs of Phasmophaga while feeding on the leaves of their
food plants.
The genus is closely related on external anatomical characters
to Hypertrophocera and Euryceromyia, less so to Euthyprosopa
1909 | Descriptions of Some New Tachinidae 245
and Pseudatractocera. The following table will serve to distin-
guish it from these forms and to point out the relationships of
each to the others:
1. Orbital bristles absent in male, front of male not over one-third head-width,
second antennal joint elongate, third antennal joint of male only a
little over twice as long as “second, arista thickened on only its basal
one-fourth to one-third, male claws elongate and female claws shorter;
facial carina present, ‘obsolete below; apical cell closed in margin
or very short-petiolate, ending well before wingtip... .Pseudatractocera
Orbital bristles in both sexes, front of both sexes never less than about
one-half of head-width and often more, second antennal joint never
elongate, third antennal joint of male alw ays greatly developed, arista
always thickened practically its whole length, claws same in both sexes
and short or but little elongate... 2
Faciala ciliate one-half way up, frontal bristles desc ending normally “below
base of antennae, discal abdominal bristles preset except in P. mert-
dionalis, no facial carina... . a)
Facialia bare, frontal bristles descending to. opposite lower border of eyes,
third antennal joint always equilateral, second aristal joint not elon-
gate, discal abdominal bristles absent, apical cell ending well before
wingtip. Sak eaten Pte a at tae ot, Wee) bo 5 ae ee © Cis ari Oe RRC Se 4
Fourth vein obsolete apically, third vein ending near wingtip; second aristal
joint elongate, third antennal joint of “male subtriangular, cheeks
HeAthye ASmWwidemaSs. eye-herclitercens ci. a1 eeneeie Euryceromyia
Fourth vein entire, apical cell very short petiolate and ending well before
wingtip; second aristal joint short, third antennal joint equilateral in
both sexes, cheeks not over one-half of eye-height....... Phasmophaga
4. No facial carina, apical cell closed in margin......... Hypertrophocera
Facial carina present, apical cell petiolate.............Euthyprosopa
bo
1S)
This study of Phasmophaga and its comparison with the above
related genera have demonstrated with considerable certainly
that Neotractocera Towns. is but the female of Hypertrophocera
Towns. In that case the latter name holds for the genus.
These genera are evidently alhed to baumhaueria, Thelymor pha,
Urophylla, and less so to Brachychaeta. Brauer and Von Ber-
genstamm put Urophylla in their group Thryptocera, but it cer-
tainly seems more closely allied with Thelymorpha. They also
refer Euryceromyia and Hypertrophocera to their group Thryp-
tocera, while they refer Neotractocera, Euthyprosopa and Pseuda-
tractocera to their group Paramacronychia along with Melano-
phrys. That the latter does not belong as referred by them is
evident from the fact that dissection of females shows the genus
to have the leaf-larviposition habit, and thus to belong to the
Echinomyiine-Hystriciine series. Hypertrophocera especially is
so clearly related to Phasmophaga in its anatomy that it seems
certain that it has the leaf-ovipositing habit. That Phasmophaga
does not belong to the Thryptocera group is clear, since the latter
genus, if we may judge from dissections of the closely allied
Bigonichaeta, must have the host-larviposition habit. Thus we
246 Annals Entomological Society of America [Vol. TI,
seem compelled to combine our genera in one group with Baum-
hauerta, etc., and if that genus proves to have the leaf-oviposition
habit this arrangement will be duly substantiated.
The European genus Pexopsis B. B (type tzbialis Meig.) seems
to form with Pseudatractocera a connection between the Baum-
haueriinae and the Masiceratinae. Like Pseudatractocera it
lacks orbital bristles in the male, but resembles the Baum-
haueriinae in the weight of its other characters.
Cyclotaphrys gen. nov.
This genus differs from Tachina (larvarum) as follows: Facialia
not at all ciliate; second aristal joint elongate; parafrontals of
male with fine hairs outside frontal row; apical cell closed in
border before wingtip; second antennal joint moderately elongate;
vibrissae not strong, inserted near oral margin.
The eyes are thinly short-hairy; parafacials moderately wide,
of even width; palpi elongate, curved, subcylindrical, a little
thickened on apical half; apical scutellar bristles suberect and
decussate; ovipositor very long and large; hind tibiae in both
sexes quite thickly short-ciliate, with two stronger bristles near
middle; fourth vein with wrinkle with bend. Claws of male
elongate. Two strong and one weak sternopleural bristles; four
postsutural bristles.
In the last-stage maggot and puparium the middle slit of
each stigma is shortened and pushed outward at lower end so as
to make the three slits appear somewhat lke an irregular circle.
The slits are sometimes very crooked and abruptly bent.
Reproductive habit, supracutaneous host-oviposition. Type,
C. anser n. Sp.
Cyclotaphrys anser n. sp.
Length 10 to 12 mm. Blackish, covered with silvery bloom.
Parafacials wholly silvery-white, parafrontals with very faint tinge of
golden in both sexes; third antennal joint three times as long as
second; palpi yellow; sides and tip of abdomen of male reddish, tip of
abdomen of female reddish.
First abdominal segment and apical borders of second and third
segments more or less shining black. The species deposits a golden-
yellow egg.
Type, No..12620, U; S.No MM.
Numerous specimens reared from importations of Euproctts
chrysorrhoea received at the Gipsy Moth Laboratory from
Simferopol, Russia (Mokchetsky).
1909 | Description of Some New Tachimidae 2447
Tachina japonica n. sp.
Length, 10 to 12 mm. _ Differs from /arvarum as follows: More
deeply colored, the golden on parafrontals heavy and extending heav-
ily on parafacials nearly to cheek grooves in male. The pollen of
mesoscutum is of a distinctly brassy shade, contrasting readily and
abruptly in both sexes with the silvery pollen of the pleurae. In
larvarum the golden shade on the parafrontals is thin in both sexes
and does not invade the parafacials to any extent, while the pollen
of mesoscutum and pleurae is concolorous and wholly silvery.
Type, No. 12621, U.S. N. M.
Many specimens of both sexes reared from importations of
Porthetria received at the Gipsy Moth Laboratory from the
vicinity of Tokyo, Japan, 1908 (Kincaid).
Paragermaria gen. nov.
Differs from Pseudogermarta as, follows:
Cheeks about one-fourth eye-height; parafacials hairy; ocellar
bristles long, reclinate; no orbital bristles in male, female with
two prochnate ones; two rows of frontal bristles on each side;
front of male wider at vertex than one.eye, face of male as wide
as both eyes; face and front of female nearly same width, except
vertex which is noticeably wider than that of male; frontal
bristles descending below insertion of arista; vibrissae strong,
decussate; second antennal joint more elongate in female than
in male, third joint more elongate in male than in female, third
joint being hardly two and one-half times as long as second in
female and more than four times as long as second in male; palpi
slender, thickened at tip; apical pair of scutellar bristles not as
strong as lateral pairs, but long and divergent. The intermediate
segments of abdomen of both sexes lack discal bristles, and first
segment is without medial marginal ones; abdomen and scutellum
clothed with suberect bristly hairs.
Reproductive habit, leaf-oviposition. Type, P. autumnalis
1135S!
Paragermaria autumnalis n. sp.
Length,9mm. _ Blackish, with more or less of gray bloom. Head
of both sexes entirely pale brassy pollinose, except the brown frontalia
and antennae, the second joint of which is reddish. Apical portion of
scutellum, whole of apical segment of abdomen of female, tip of apical
segment of male, and tibiae reddish. Pleurae, scutellum, femora,
and all of abdomen except basal segment and more or less of posterior
margins of other segments with a silvery bloom showing less of brassy
if any than that of head. Mesoscutum with less of pollen, showing
four heavy black vittae and a median fifth one behind suture.
248 Annals Entomological Society of America [Vol at;
Type, No. 12622, U.S. N. M.
Numerous specimens of both sexes taken through September,
1908, Melrose Highlands, Massachusetts, on flowers of Aster sp.,
also on Solidago flowers and foliage. The dissected females
uniformly contained a long coiled uterus filled with black micro-
scopic eggs ready for leaf-deposition.
Sisyropa hemerocampae n. sp.
Syn., Exorista griseomicans Coq. (non. v. d. W.) pt.
Length, 7 to8 mm. Front about or nearly width of one eye in
female; more narrowed, especially at vertex, in male. Parafrontals,
parafacials and facial plate entirely silvery; frontalia brown; antennae
brownish with slight silvery bloom, articulations shghtly fulvous;
palpi yellowish. Thorax and scutellum yellowish, dusted with silvery,
with five vittae. Pleurae silvery. Apical scutellar bristles long and
strongly cruciate. Abdomen blackish, sides of second and third seg-
ments above broadly reddish, whole ventral surface yellowish-red
except median vittae and anal segments. Legs reddish-yellow, tarsi
brownish. Tegulae whitish, wings clear.
Type, No. 12623, U.S. N. M.
Two females labeled: Gipsy Moth Laboratory, No. 820-0,
1907. One male, labeled: Washington, D. C., F. D. Couden,
Coll.- Adult issued October 2, 1907. All three specimens reared
from Hemerocampa leucostigma.
This species has been collected in the field at Melrose High-
lands, Massachusetts, the female specimen bearing Gipsy Moth
Laboratory No. 1976 being indistinguishable from the above
specimens.
The genus Szsyropa, which belongs in the subfamily Hem1-
masiceratinae, may be distinguished from the closely similar
genus Eumasicera (described below) of the Masiceratinae as
follows: Szsyropa has_the face and front of female very narrow,
nearly equilateral, and equal to width of one eye; cheeks and
parafacials extremely narrow, the latter at narrowest equal to
half length of second antennal joint or even less; eyes thickly to
densely pilose; palpi moderately stout basally, a httle enlarged
at tip; scutellum and abdomen with thickly placed erect hairs;
apical pair of scutellar bristles long; antennae inserted well above
eye-middle. Reproductive habit, larviposition.
1909] Descriptions of Some New Tachinidae 249
Eumasicera gen. nov.
Differs from Sizsyropa (as above outlined) as follows: Face
and front of female wider than one eye; cheeks and parafacials
wider, latter more rapidly widening above; eyes thinly pubescent ;
palpi slender basally and abruptly widened and _ thickened
apically; scutellum with sparse erect hairs, abdomen with short
appressed bristles; apical pair of scutellar bristles short; antennae
inserted about on eye-middle.
Reproductive habit, leaf oviposition. Type L. coccidella n. sp.
Eumasicera coccidella n. sp.
Length,8 mm. Faintly brassy pollinose all over, except the face
which is silvery, and the first abdominal segment and legs which are
blackish. Palpi reddish-yellow.
Type, No. 12624, U.S. N. M.
One female, No. 1975, Gipsy Laboratory.
The minute eggs of this species, obtained from the tubular
coiled uterus of the female, are black and show a remarkable
light-colored subdentate fringe around the edge, the indentations
marking divisions in the fringe and the divisions exhibiting a
network structure shot with microscopic holes. The chorion of
the egg is reticulate after the ordinary plan. These eggs are
seen to be flattened and the chitinous chorion marks the superior
surface, while the surface attached to the leaf at time of ovi-
position is furnished with a thin membraneous unchitinized
covering.
Rileyella gen. nov.
Differs from Achaetoneura as follows: Both sexes without
median macrochaetae on first and second abdominal segments
above, either discal or marginal.
Type, Tachina aletrae Riley.
Eucelatoria gen. nov.
This genus is erected for Tachina (Masicera [Frontina])
armigera Coqt.
Orbital bristles present in female, absent in male. Eyes bare.
Abdomen with discal bristles on intermediate segments in male,
but without median bristles either marginal or discal on first two
segments in female. Median ventral carina present in female,
strongly developed, piercing sheath present.
Reproductive habit, subcutaneous host-larviposition.
Annals Entomological Soctety of America [Wolds
to
On
O
Cordyligaster septentrionalis n. sp.
Length, 11 to 13 mm. Ground color black, more or less shining;
head silvery-white pollinose, thinner at vertex both before and behind;
frontalia brown or black, somewhat shining; antennae black, thinly
silvery dusted; pleurae, coxae and narrow bases of last three abdomi-
nal segments silvery-white; mesoscutum and scutellum shining-metal-
lic, without vittae; tegulae transparent; wings with wide costal border
smoky-yellowish, faintly tinging inner portion. Apical cross-vein
deeply bowed inward, forming right-angle with fourth vein. Palpi
pale-yellowish but appearing blackish from black hairs and silvery
bloom. Femora very thinly dusted with silver.
ype, Nov 1262'5.sUo IN: iM.
Numerous specimens of both sexes from Plummers Island,
Maryland, June and July, 1907 (Dr. A. K. Fisher).
Parachaeta Codt.
The generic description of Parachaeta needs revision. Only
the male was described. That sex is without macrochaetae on
the second abdominal segment, while the female has a pair of
stout macrochaetae on the hind margin of that segment and
sometimes a shorter pair between them. The absence of macro-
chaetae on the second segment thus becomes a secondary sexual
character in this genus.
The common New England species is the Parachaeta bicolor
of Coquillett’s Revision but not of Macquart. The female is
darker in coloration than the male, and shows no reddish on sides
of abdomen in the specimens that I have examined. « The
species does not agree well with Bigot’s description of tmermis.
The type of the latter must be examined.
NEW FOSSIL INSECTS FROM FLORISSANT, COLORADO.
By T. D. A. COCKERELL.
It was not at first intended to do any work at Florissant dur-
ing the summer of 1909, but two of my students, Messrs. Rusk
and Duce, having become interested in the study of fossils,
arranged to go there for a short time. Their collections include
the usual percentage of insects, several of which are new and are
herewith described.
HYMENOPTERA.
Eriocampoides revelatus sp. n. (Tenthredinide; Phyllotomine.)
Length about 12 mm.; anterior wing about 84; thorax black; legs
hardly visible, but apparently black; abdomen reddish-brown, very
robust, 44 mm. broad; wings ample, faintly dusky, anterior wings
with stigma and nervures brown.
Venation not essentially different from that of Eriocampoides
(cf. Macgillivray, Pr. U.S. Nat. Mus., 1906, pl. XXX, figs. 52 and 53);
the most striking peculiarity is in the hind wings, in which the vein
which should form the truncate apex of the radio-medial cell (marked
M in Macgillivray’s figures) is wanting, the discal cell being actually
joined to (contiguous with) the radial for a very short distance, and
the end of the radiomedial cell. consequently pointed. In other
respects, as the form of the lanceolate, cubital and discal cells, the
hind wing agrees with that of E. e@thiops. The peculiarity referred
to may be a variable one, as on one side the medio-cubital vein is
broken and a little displaced, but looks as if it had been so attached as
to leave an extremely short interval between the discal and radio-
medial cells; in this case the description should read, discal cell touch-
ing or almost touching the radial, which would afford a very good
distinction from the living species. The upper side of the discal cell
is about 629 y long.
The following measurements of the anterior wings are in microns:
Cross-nervure of marginal cell, which is long and curved but not
bent, about 1105.
First submarginal cell on first discoidal 527.
Second submarginal cell on marginal 1020, on first discoidal 510,
on third discoidal 884.
Third submarginal cell on third discoidal 340.
Second transverso-cubital nervure to insertion of marginal cross-
nervure 1020; the latter to third transverse-cubital 306, and to apex
of marginal cell 1496.
Basal side of first discoidal cell (basal nervure) 1275, its apical
side 765.
Lower end of basal nervure to transverso-medial 680, length of
the latter 697.
Second discoidal cell on first 782.
Beginning of cross-nervure of lanceolate cell to lower end of trans-
verso-medial 1054.
251
252 Annals Entomological Society of America [Vol. EH,
The insect resembles EF. ethiops in the distance of the transverso-
medial from the basal, the basal without a bend, and the first trans-
verso-cubital so placed that if prolonged it would form a very acute
angle with the costa. It resembles FE. varipes in the long (longer than
in varipes) and very oblique marginal cross-nervure, joining the third
submarginal cell near its end. In the bend of the nervure bounding
the lanceolate cell below, it is intermediate between @thiops and
varipes. In the long pointed marginal cell it is nearest to var1pes, but
in the comparatively slight divergence of the two sides. of the first
discoidal it resembles ethiops. It is the first of its genus to be found
fossil.
Habitat.—Miocene shales of Florissant, Colorado (Wadallard
Rusk, 1909). This has the costal cross-nervure, and is an
Eriocampoides as defined by Macgillivray.
At Station 13 B, Mr. Duce found a very beautiful specimen of
Megaxyela petrefacta Brues.
Odynerus terryi sp. nov. (Eumenidz.)
Length probably about 18 mm., but only the basal-part of the
abdomen is preserved; head and thorax black, densely punctured;
head about 4 mm. long and 3? mm. broad. The structural details
cannot be made out, except that the ocelli are large (255 mw diam.),
the scape is slender, and the mandibles are sharply dentate; the an-
tennal sockets are about 544 » apart; thorax about 6 mm. long and
5 wide, the large prothoracic lobes as usual in the genus; posterior
lobes of the metathorax, on each side of the base of the abdomen, very
large and conspicuous, being about 850 long and 765 wide, densely
punctured; first abdominal segment broad and short, formed and
colored, so far as can be seen, precisely as in O. diffinis Sauss. (Bing-
ham, Hymenop. British India, I, f. 106) except that it is shorter, and
therefore broader in proportion to its length,—it may, however, be
shaped precisely as in dijfinis, 1f the clear band, which seems to bound
the segment, really traverses its disc, the whole of the structure,
which looks like two segments, being really only one. (After repeated
examination, I feel convinced that the latter explanation is correct.)
Anterior wings about 124 mm. long, folded as usual in the group,
dark fuliginous; stigma reddish, nervures piceous. Venation normal
for Odynerus; marginal cell at end very obliquely truncate, ending in
a point away from costa; basal nervure with its upper end remote
(about 850 «) from stigma; cubital nervure not bent at end of first
discoidal cell; third transverso-cubital nervure arched inwards; both
recurrent nervures joining second submarginal cell, the first about 425 yu
from its base, the second about 289 from apex, the distance between
them about 544. The second submarginal is greatly narrowed above
its width on marginal being only about 170 ». Depth of marginal
cell 986 yp.
Hind wings pallid, with the anal and transverso-medial (or trans-
verse anal) nervures scarcely forming any angle or bend at their junc-
1909 | New Fossil Insects from Florissant, Colorado 253
tion; a character very distinctive of the Odynerines; in the Vespines
they form a strong angle, and the anal is usually continued to the
margin.
Habitat—Miocene shales of Florissant, Colorado, tg09 (Terry
Duce). <A fine species, easily known from the Odyneri previously
described from Florissant by its large size.
Messrs. Duce and Rusk obtained a single anterior wing, well
preserved, of Megachile predicta Ckll. Can. Ent. 1908, p. 31.
This shows that the wing is hyaline, faintly reddish, with the
nervures and stigma pale ferruginous.
DIPTERA.
Syrphus aphidopsidis n. sp. (Syrphidz.)
Q (apparently); length 144 mm., but owing to pressure, the
pointed apex of the abdomen, usually retracted in cabinet specimens,
is extended, giving the insect a rather peculiar appearance, and adding
to the length. Width of head about 3 mm., of thorax 3}, of abdomen
‘about 4, length of wings 9. Eyes bare, about 900 » apart on vertex,
the facets of uniform size, about 34 » diameter; antenne normal, the
apical joint large and rounded, with the usual stout bristle; thorax
above coal-black, without light markings; abdomen with very broad
light bands on the first three segments, but all interrupted in the middle
to the extent of about a third of amm.; on the first segment the base
is dark, but the rest light except a large median inversely cup-shaped
area about 24 mm. in diameter, its rim resting on the hind margin,
its other end connected by a stem with the dark basal area; second
segment with a broad basal band, interrupted in the middle, straight
behind, except at extreme sides where it is produced posteriorly;
third segment with the pale areas widening from the interruption, so
that they are triangular in form; fourth segment with a fine continuous
pale line at base, widening laterally; wings faintly dusky; hind femora
stout, about 680 » diameter, their tibia very slender.
Venation compared with that of Syrphus arcuatus, Eupeodes
volucris and Sceéva pyrastrt; measurements in microns:
(1) First vein normal; after reaching the margin it really con-
tinues, as in the living forms, to the end of the second, so that the
marginal cell, while apparently open, is in reality closed.
(2) Second vein arched, and at its tip deflected upwards; this
agrees much better with Syrphus than with Eupeodes or Sceva.
(3) Third vein beyond the anterior crdss-vein straight, with a
just observable arching; this is quite different from Sce@va; also differ-
ent from S. arcuatus and Eupeodes, in that what arching there is
begins at the cross-vein, instead of being confined to the distal part.
(4) Spurious vein very distinct, bisecting the cross-vein a little
higher up than in Sceéva.
(5) Anterior cross-vein not at all oblique (more or less oblique
in the living species), placed about 1020 from the base of the discal
cell, and of course much before the middle.
254 Annals Entomological Society of America [Volo
(6) Base of discal cell with a diameter of 425; apex rounded
and quite remote (a distance as great as the length of the anterior
cross-vein) from the lower corner of the first posterior cell. This is
unlike the three recent species compared, but the character occurs, even
more pronounced, in Brachyopa notata and other Syrphide. The outer
side of the first posterior shows a very strong double curve, and the
apex of the cell is quite remote from the margin; all this essentially
as in Baccha dimidzata.
(7) Cross-vein from anal to discal cells about 595 long, thus con-
siderably longer than base of discal, which is not at all the case in the
recent forms compared. This condition exists in Xylota pigra, but
usually when there is a difference, it is the other way among the Ameri-
can Syrphide.
(8S) Anal cell normal, closed some distance from margin of wing.
Habitat.—Miocene shales at Florissant, Colorado, 1909 (W1I-
lard Rusk). The specific name suggests that it may have fed on
the genus of Aphididae, most abundant in the vicinity. The
insect seems not to be quite typical Syrphus, but it appears to go
better there than elsewhere. Easily known from S. wuillistont
Ckll., also fossil at Florissant, by its larger size.
Dioctria (?) florissantina (CkIl.) (Asilidaee; Dasypogonine).
When describing Leptis florissantina (Am. Jn. Sci., Jan. 1909,
p. 58), I did not attach enough importance to the length of the
second vein. Renewed study shows that the venation is exactly
that of the Asilid genus Dioctria, except that the cell in the forks
of the third vein does not widen apically. The abdomen also
agrees with Dioctria, but the thorax is much too broad and round-
ed in front, and the head and eyes, while ordinary for a Leptid,
do not look like those of an Asilid. It is possible, however, that
they are somewhat distorted.
Microstylum (?) destructum sp. nov. (Asilide; Dasy pogonine.)
This insect appears to represent a new genus, but by a great
misfortune the pick fell so as to shatter and destroy the whole of
the apical half or more of the wings and abdomen, and it is not
desirable to attempt a generic diagnosis from what remains.
Body exceedingly robust, head and thorax shining coal-black, the
latter about 6 mm. long; structure of head not ascertainable; abdo-
men reddish-brown with the sutures hyaline, 63 mm. broad at base;
legs slender, dark, hind femur 765 » broad near end, without bristles;
hind tibia about 544 ” broad, with very. small inconspicuous bristles.
Venation agreeing with Muicrostylum flaviventre, so far as pre-
served, except as follows:
1909] New Fossil Insects from Florissant, Colorado 255
(1) Stem of second and third veins (from first to divarication)
only about 1020 » long, and leaving first at a somewhat larger angle;
fork much wider, so that at about 2800 # from the fork the second and
third veins are about 1055 yu apart.
(2) For the length of 2800 the third vein, very clearly visible,
shows no sign of the cross-vein to the fourth, but at this point (where
the break occurs) the fourth begins rapidly to ascend, showing that
the cross-vein is very little beyond. It results from this that the first
basal cell is extremely long, a character of the Mydaide.
(3) The fourth vein before the discal cell is not in a straight line
with the part bounding the upper side of the cell, there being a very
decided bend.
(4) The junction of the discal and second basal is longer, about
1870 uw. (The cross formed at the lower basal corner of the discal cell,
and the adjacent parts toward the anal, are exactly as in Microstylum.)
(5) The anal cell is narrowly (170 ) open.
(6) The fifth posterior cell has its upper apical corner cut off by
the deflection (to reach the margin at an acute angle) of the terminal
section of the fifth vein. This condition is approached in Mydaide,
and slightly in some Apioceride.
(7) The vein which bounds the outer (marginal) side of the
closed fourth posterior cell, which in the Asiline is directed upwards
from the lower corner of the cell, but in Microstylum is approximately
parallel with the margin, is actually directed, though very obliquely,
toward the wing-margin, being broken (through the fracture of the
specimen) after a course of 800 w, when 170 » from the margin, its
beginning being about 510 w from the margin. I cannot find any genus
showing this character, or any nearer approach to it than Microstylum .
Habitat.—Miocene shales of Florissant, Colorado, 1909 (Wul-
lard Rusk). A true Microstylum (M. wheeleri Ckll.) has been
described from the Florissant shales. In addition to the original
type, we have now also a specimen found by Mr. Geo. N. Rohwer
in 1907 at Station 14.
Nicocles miocenicus n. sp. (Asilidee; Dasypogonine.)
Length 133 mm.; wings 9; length of abdomen 91; its breadth
23/,; hind femur about 4 mm.; black, rather robust, the abdomen
cylindrical, formed as in NV. emulatar Loew; wings faintly dusky, not
appreciably spotted; legs robust, including the tarsal joints; hind tibia
much longer than its femur; hind basitarsus about 1440 » long and
580 broad; thorax and abdomen without visible hairs or bristles;
femora not visibly hairy; face with some bristles. In the following
account of the venation, comparison is made with Williston’s figure
of Nacocles rufus (N. Am. Dipt., p. 194) and Verrall’s of Dasypogon
teutonus (British Flies, v. p. 708).
Auxiliary and first veins normal, thickened, their terminations
about 2720 (this and the following measurements in microns) apart. °
Second vein with a gentle upward curve at the end, not so strong
as in Dasypogon.
256 Annals Entomological Society of Amzrica [Vole ie
Second submarginal cell long and slender; its length about 2620;
its breadth in middle 374, and near end about 510; the upper branch
of the third vein more bent at the fork than the lower. In Dasypogon
this cell is very much wider apically, and in Nicocles rufus considerably
sO.
Anterior cross-vein 1310 from the fork of third vein and 850 from
end of discal cell; in being beyond middle of discal cell it differs con-
spicuously from Dasypogon.
Second and third posterior cells normal; first posterior not widen-
ing apically. (It widens in N. rufus.)
Fourth and fifth posterior cells normal, widely open.
Anal cell open, about as in Dasypogon. (Closed in N. rufus.)
_ This seems to be a Nicocles, a genus well represented in America °
today. The type of Nicocles, according to Verrall, has the anal cell
open.
Habitat.—Miocene shales of Florissant, Colorado, 1909 (Terry
Duce). While on Asilide I wish to record the occurrence of
Laphria fernaldi (Dasyllis fernaldt Back) at Teller Lake (near
Tolland), Colorado, alt. 9575 feet (J. Henderson). A single male
was taken July 29, 1909. This fine species, described vaguely
from ‘‘Colorado,’’ curiously simulates the red-banded Bombi
common in the same vicinity. This resemblance to Bombus
has been noticed by P. R. Jones" (Trans. Am. Ent. Soc., 1907,
p. 282.)
» EXPLANATION OF PLATE XXVIII.
Fig. 1. Eriocampoides revelatus Ckll., venation.
Fig. 2. Microstylum destructum, venation.
Fig. 3. Megaxyele petrefacta Brues.
Fig. 4. Syrphus aphidopsidis Ckll.
Fig. 5. Odynerus terryi CkIl.
ANNALS E. S. A.
Radial.
lanceolate
or anal,
hind wing
Ericcambeidses rivelatus.
Fig. 1.
Fig. 4.
T. D. A. Cockerell.
Vor. II, PLATE XXVIII.
: 4
7 ae
173) 3
=a ee |
oa
A SYNOPSIS OF THE AMERICAN SPECIES OF
SCOLIONEURINAE.
By Arex. D. MacGILiivreay.
The European species of this subfamily were originally
referred to the genus blennocampa. konow in 1886 removed one
of the species from blennocampa under the generic name of
Entodecta and in 1890 the remaining species under the name of
Scolioneura. These genera, he has associated with blennocampa |
and its allies under the tribal name of Blennocampides! in his last
analysis of the group. The first American species was described
by Norton in 1867 as Selandria capitalis, later, 1895, referred to
Scolioneura by Marlatt? and two new species described. 5S. A.
Forbes* in 1885 described Metallus rubs from two male specimens
bred from larvae mining in the leaves of cultivated blackberries.
This species was referred to the genus Fenusa by Cresson‘ in
1887 as a possible variety of Fenusa curta Nort. Konow in the
Genera Insectorum, evidently on the authority of Cresson, makes
Metallus a synonym of Fenusa and adopts rubi as a good species.
An examination of Forbes’s type proves it to belong to the Scol-
ioneurinae. Since Metallus antedates both Entodecta and Scolto-
neura, if it should be found later that only two generic names
can be retained in this subfamily, Metallus will have to be
used for one of them. A fifth species was added by Kincaid* in
tgoo from Alaska under the name of Fenusa alaskana, a species of
FEntodecta. Finally a sixth species was described by Konow’,
Entodecta humilis, from Alaska in 1908. The present writer in
19067 separated Scolioneura and Entodecta from the Blennocam-
pides as a distinct subfamily, based on the divergent condition
1. Konow, F. R. W.—Genera Insectorum. Hymenoptera, Family Ten-
thredinidae. Fascicule 29, 1905, 76-90. ;
2. Marlatt, C. L.—The American species of Scolioneura Knw. Proc.
Ant. Soc. Wash., III, 1895, 234-236.
3. Forbes, S. A.—Fourteenth report of the state entomologist on the
noxious and beneficial insects of the state of Illinois. Springfield, 1885, p. 87,
joll, IDSs sale Tf
4. Cresson, E. T.—Synopsis of the families and genera of the Hymenop-
tera of America, north of Mexico. Philadelphia, 1887, p. 160.
5. Kincaid, Trevor. Papers from the Harriman Alaskan expedition.
VII. The Tenthredinoidea. Proc. Wash. Acad. Sci. II, 1900, 345.
6. Konow, F. R. W.—Zeit. System. Hymen. Dipter, VIII, 1908, 84-85.
7. MacGillivray, A. D.—A study of the wings of the Tenthredinoidea, a
superfamily of Hymenoptera. Proc. U. S. Nat. Mus., XXIX, 1906, 649.
259
260 Annals Entomological Society of America [Vol. II,
of the medio-cubital cross-vein and the free part of M,,, in the
front wings and from the Fenusinae, another small group of
genera included in the Blennocampides by Konow, by the form
of the cell R,,, of the hind wings. The cell R,,, being closed
throughout and appendiculate at apex and with vein R, reaching
the margin of the wing distinctly before,the apex in the Scolio-
neurinae and the cell R,,, rounded at apex and open and the
vein R, reaching the margin of the wing at or beyond the apex
in the Fenusinae.
Konow® describes the larvae of five species, all are leaf-
miners, two are found on Betula, one on Tilia, one on Geum, and
one on Rubus. In addition to the record of Forbes of Metallus
rubt as a leaf-miner on Rubus as given above, Houghton® has.
reported a species on Rubus from Delaware of which specimens
were named by the writer as Scolioneura capitalis Nort.
This subfamily can be differentiated from its congeners as.
follows:
SCOLIONEURINAE.
Head transverse; the clypeus long, never deeply emarginate;
the antennae with nine segments; the eyes oval in outline, the
inner margin straight or uniformly convex, adjacent to the base
of the mandibles; front wings with the radial cross-vein, the
radio-medial cross-vein usually, and the free part of R, and R;
present; the medio-cubital cross-vein joined to Sc,+R+M at or
near the origin of the free part of media; the medio-cubital cross-
vein and the free part of M,,, strongly divergent behind; the
free part of M,+Cu, arising at or near the middle of the cell
M,; the free part of media and the radial sector straight or
gently curved near their origin, not strongly bowed; the medio-
cubital cross-vein strongly bowed and joining cubitus about
midway between the origin of cubitus and the point of separation
of M, and M,; the hind wings with the cell R,,, closed and ap-
pendiculate at apex, very rarely open; the vein R, reaching the
margin of the wing distinctly before its apex; the free part of
R, and the transverse part of M, wanting; the second anal cell
distinctly petiolate; the claws appendiculately toothed.
8. Konow, F. R. W.—Systematische Zusammenstellung der bisher be-
kannt gewordenen Chalastogastra. Zeit. Syst. Hymen. Dipter. I, 1901, 228-229.
9. Houghton, C. O.—The Blackberry Leaf-miner. Entom. News, XIX,
1908, 212-216.
T909| American Species of Scolioenurinae 201
GENERA OF SCOLIONEURINAE.
a. Front wings with the radial cross-vein and the free part of R, interstitial;
the antennal furrows practically continuous throughout........
Scolioneura Knw.
aa. Front wings with the radial cross-vein and the free part of R, widely sepa-
rated; the antennal furrows broadly interrupted on the middle of the
front.
b. Antennae with the second segment distinctly longer than broad; the
first and second segments together as long as or longer than the
Haulbigo tei sro crochet Sic cg 2.6 canon eee ie Caen Parabates n. gen.
bb. Antennae with the second segment annular, broader than long; the
first and second segments together shorter than the third.
c. Eyes with their inner margins straight and parallel, not converg-
ing below; the front distinctly wider than high........
Polybates n. gen.
ec. Eyes with their inner margins uniformly convex and converging
below; the front not as wide as high.
d. Front wings with the free part of R, and the radial cross-vein
inclimed at ‘ditterent angles. ....4..: =. Metallus Forbes
dd. Front wings with the free part of R, and the radial cross-vein
inclined at approximately the same angle........
Entodecta Knw.
Scolioneura Knw.
Head transverse, the front distinctly broader than high;
the inner margin of the eyes straight and parallel, the eyes
not converging below; the antennal furrows deep and distinct,
practically continuous from the antennal fovea to the occiput;
the clypeus squarely truncate, distinctly more than half as wide
at apex as at base; the antennae long and filliform, the second
segment shorter than the first, broader than long, globular, not
annular, the first and second segments together shorter than the
third; front wings with the free part of R, and the radial cross-
vein interstitial and inclined at the same angle; the free part of
M,,Cu, received at or beyond the middle of the cell rst A+
2nd 2nd A; media strongly angular at the point of separation
of M,,,and M,,,. Type Tenthredo betulets Kg.
This generic name as here used is restricted to those species
in which the radial cross-vein and the free part of R, is interstitial.
The generic diagnosis is based on a specimen of the type species
identified by Dr. Fr. W. Konow. For a further discussion of
the characters used for this genus see under Entodecta.
Scolioneura populi Marl.
Body whitish with the following parts black: the antennal fur-
rows, the antennal fovea, a spot around the anterior ocellus, a trans-
verse band on the postocular area, the antennae beyond the first seg-
ment, a spot on the median and lateral lobes of the mesonotum, the
scutellum, the metathorax, the apex of the pectus, the abdomen above,
262 Annals Entomological Society of America [Vol wile
and the saw-guides; antennae not thickened at middle, the first and
second segments globular, subequal in length, together five-sixths the
length of the third, the third as long as the fourth and fifth together,
the fourth shghtly longer than the fifth, the fifth and sixth subequal
and each slightly longer than the seventh, eighth and ninth, which
are subequal, the ninth with parallel sides and a bluntly rounded
point; the clypeal foveae deep and prominent; the supraclypeal* area
slightly convex and not prominent; the antennal furrows deep, promi-
nent, extending from the clypeal fovea to the occiput, and almost
straight; the median fovea an inverted wedge-shaped depression, long-
er than broad; the postocullar area convex, well separated; the inter-
ocellar furrow distinct, continuous, straight; the ocellar basin fan-
shaped, connected with the interocellar furrow; the posterior meta-
tarsus shorter than all the following segments together; the second
and third segments of the tarsi subequal; the front wings with the free
part of R, twice as long as the free part of R;; the stigma convex below,
about three times as long as broad, angled at the point of attachment
of the radial cross-vein; the veins, including the costa and the stigma,
luteous; the saw-guides moderately robust, truncately rounded, and
somewhat pointed at the upper apical angle. Length 4 mm.
Habitat.—Las Cruces, New Mexico. - Described from two
females received from Prot te D. A.Cockerell:
The male according to Marlatt does not differ except in
having a greater percentage of white or brownish white on the
body.
Parabates n. gen.
Head transverse, front not as broad as high; the inner mar-
gin of the eyes uniformly convex, the eyes distinctly converging
below; the antennal furrows obsolete except adjacent to the
antennal foveae and behind the lateral ocelli; the clypeus squarely
truncate, distinctly more than half as wide at apex as at base;-
the antennae long and filiform, the second segment as long as
the first, longer than broad, as wide as the first segment, not
annular, the first and second segments together as long as or
longer than the third; the front wings with the free part of R,
and the radial cross-vein not interstitial and inclined at the
same angle; the free part of M,+Cu, received at the middle of
the cell rst A+ 2nd 2nd A; media strongly angular at the point of
separation of M,,, and M,,,; the radial sector strongly bowed at
base. Type Parabates listrionicus n. sp.
* Marlatt in his monograph of the Nematinae uses the term hypoclypeal
plate for this area and the writer has referred to it thus far as the hypoclypeal
area. The word hypoclypeal is a misnomer, meaning below instead of above
the clypeus, it is also a hybrid in origin and should be discarded.
1909] American Species of Scolioneurinae 263
Species of Parabates.
a. Vertical portion of the antennal furrows punctiform, only slightly longer
RAMS a-rohs <i) So. A eas eet eee Pat's histrionicus n. sp.
aa. Vertical portion of the antennal furrow not punctiform, extending from the
Occiptimto beyond) ther lateralmocellitiys. 2-0-6 .2 4 ete inspiratus n. sp.
Parabates histrionicus n. sp.
Body black with the knees, the front and middle tibiae and their
tarsi, and the posterior tibiae and metatarsi, white or infuscated;
the antennal furrows deep above and below, interrupted on the middle
of the front, the lower ends connecting with a broad, deep antennal
fovea, extending to opposite the middle of the median fovea, inter-
rupted by a transverse ridge not completely filling the furrow, the
furrow punctiform above the ridge, longer than broad, the vertical
portion punctiform, not extending beyond the lateral ocelli, slightly
longer than wide; the postocular area very wide and very short, de-
pressed below the area in front of the interocellar furrow; the interocel-
lar furrow faint; the middle of the front uniformly convex, the ocellar
basin practically wanting, a slight depression around the median
ocellus and connected with the interocellar furrow by a very fine
furrow; the median fovea deep, with broadly inclined sides, longer than
broad; the supraclypeal area strongly elevated between the antennae,
ridge-like, uniformly convex below the median fovea; the antennae
slender, the first and second segments globular, subequal in length,
together slightly longer than the third segment, the third as long as
the fourth and fifth together, the fourth and following segments sub-
equal in length, the ninth segment gradually narrowed toward the
apex, pointed on one side, the fourth and following segments slightly
serrate on one side, all the segments densely hairy; the clypeus trun-
cate at apex; the wings slightly infuscated, the veins and the costa
black, the stigma except on its front margin brownish, a fuscous band
between the tip of the third anal vein and the base of the cell Ist A+
2nd. 2nd. A.; front wings with the free part of R, three times as long
as the free part of R;; the stigma broadly rounded below, truncate at
apex, twice as long as wide, broadest before the middle; the posterior
metatarsus shorter than all the following segments together, the sec-
ond segment twice as long as the third; the saw-guides straight on the
upper margin, convex below, obliquely, convexly truncated at apex,
with a blunt point at its upper angle, the surface polished, the apex
and the lower margin fringed with setae longer than the width of the
saw-guides. Length 3 mm.
Habitat.—Olympia, Washington (Trevor Kincaid); Colorado
(C. F. Baker); and Nevada. Type in the collections of Cornell
University and paratypes in the collections of the U. S. National
Museum.
264 Annals Entomological Society of America [Vol. II,
Parabates inspiratus n. sp.
Body black with the knees, the tibiae, and the tarsi infuscated;
the antennal furrows deep and broad, above and below, interrupted on
the middle of the front, the lower ends connecting with the antennal
fovea, transversely divided by a ridge just above the base of the
antennae, the portion above the ridge punctiform, deep and broad,
the vertical portion deep, extending from the occiput to beyond the
lateral ocelli, much longer than wide, the interocellar furrow distinct;
the ocellar basin depressed around the median ocellus, with a broad
furrow extending from this basin to the interocellar furrow; the median
fovea broad, rather shallow, narrower than the space between the
antennae; the supraclypeal area convex and prominent; the antennae
slender, the first and second segments globular, subequal in length,
together subequal in length to the third, the third twice as long as the
fourth, the fourth and following segments subequalin length, all the
segments densely hairy; the wings slightly infuscated, the veins, includ-
ing the costa and the stigma, brownish, not with a fuscous band be-
tween the tip of the third anal vein and the base of the cell Ist A+
2nd. 2nd A; the posterior metatarsus subequal in length to the four
following segments; the saw-guides convex above and below, rounded
at apex below and forming an evenly rounded point above, the entire
surface coarsely and closely punctured, covered with fine, short setae.
Length 4 mm.
Habitat.—Nevada. Type in the collections of the U. S.
National Museum.
Polybates n. gen.
Head transverse, the front distinctly broader than high;
the inner margin of the eyes straight and parallel, the eyes not
converging below; the antennal furrows obsolete except behind
the lateral ocelli; the clypeus slightly, broadly emarginate,
distinctly more than half as wide at apex as at base; the antennae
long and filiform, the second segment much shorter than the
first, annular, not as wide as the first, broader than long, the
first and second segments together shorter than the third; the
front wings with the free part of R, and the radial cross-vein
not interstitial, inclined at the same angle; the free part of
M,+Cu, received distinctly before the middle of the cell rst
A+end 2nd A; media gently curved at the point of separation of
M,,,and M,,,; the radial sector almost straight, slightly oblique,
at base. Type Polybates slossonae n. sp.
1909] _ American Species of Scolioneurinae 265
x
Polybates slossonae n. sp.
Body black with all the legs entirely white; the antennal furrows
punctiform below, longer than wide, hardly more than a prolongation
of the dorsal margin of the antennal fovea, the vertical portion of the
antennal furrows punctiform behind the lateral ocelli and extending
below the lateral ocelli for about the width of an ocellus; the post-
ocellar area flat; the interocellar furrow distinct, broadly concave be-
hind; the ocellar basin strongly convex, very slightly depressed in
front of the median ocellus; the median fovea deep, punctiform, as
broad as long, situated just above and between the bases of the anten-
nae; the supraclypeal area uniformly, strongly, convexly elevated; the
antennae slender, the second segment much shorter than the first,
annular, the first and second segments together less than half the
length of the third, the third segment very slightly longer than the
fourth, the fourth, fifth and sixth subequal, the seventh, eighth, and
ninth subequal and slightly shorter than the sixth, the ninth with
straight sides uniformly rounded to a point at the middle of the apex,
all the segments hairy; the wings hyaline, the veins, including the
costa and the stigma, brownish; the front wings with the free part of
R, about twice as long as the free part of R,; the stigma over twice as
long as wide, its hind margin uniformly, convexly rounded from base
to apex; the posterior metatarsus three-fifths the length of all the
following segments together, the second segment slightly longer than
the third and one-third the length of the metatarsus; the saw-guides
straight above and below, converging to a bluntly rounded point at
2
apex. Length 3 mm.
Habitat.—Franconia, New Hampshire. Named for its col-
lector, Mrs. Annie Trumbull Slosson. Type in the collections of
Cornell University and paratype in the collections of the U. S.
National Museum.
Metallus Forbes.
Head transverse, the front not as broad as high; the inner
margin of the eyes uniformly convex, the eyes distinctly converg-
ing below; the clypeus squarely truncate, distinctly more than
half as wide at apex as at base; the antennal furrows obsolete
except adjacent to the antennal fovea and behind the lateral
ocelli; the antennae long and filiform, the second segment dis-
tinetly shorter than the first, wider than long, more or less annu-
lar, the first and second segments together distinctly shorter
than the third; the front wings with the free part of R, and the
radial cross-vein not interstitial and distinctly inclined at differ-
ent angles; the free part of M,+Cu, received distinctly before
the middle of the cell rst A+ 2nd 2nd A; media gently curved at
the point of separation of M,,,and M,,,; the radial sector almost
straight at base. Type, Metallus rubs Forbes.
266 Annals Entomological Society of America Vola
The generic name Metallus has been practically disregarded
since its publication in 1885. It is the earliest proposed name in
the subfamily and will therefore take precedence over all others.
Species of Metallus.
a. Stigma three times as long as broad; the front punctured around the base
of they janitenmae ? Ay: bt ec Beto ee en cusey tae ees canadensis Marl.
aa. Stigma twice as long as broad; the front not punctured around the base ~
of the antennae.
b. Front wings with the free part of M,+Cu, joining the cell M, distinctly
beyond, thesmiddilestes, 95 0%, teeeals fie eee eee capitalis Nort.
bb. Front wings with the free part of M,+Cu_, joining the cell M, at middle.
c. Mesonotum and scutellum rufous............... rohweri n. sp.
cc. Mesonotum and scutellum black................. rubi Forbes
Metallus canadensis Marl.
Body shining black with the median and lateral lobes of the meso-
notum rufous and the labrum, clypeus, mandibles, antennae, angles
of the pronotum, tegulae, four anterior legs below the bases of the
coxae, and the trochanters, tibiae, and tarsi of the hind legs, orange-
white; the head punctured around the base of the antennae, with a
faint transverse furrow over the antennae; the antennal furrows not
nearly complete; the lateral walls of the ocellar basin sharply defined
but low; the ocellar basin well defined above and connected with the
straight interocellar furrow, the interocellar furrow strongly con-
tinuous; antennae very short and stout, with the third segment not
quite as long as the fourth and fifth together, the second segment as
long as the first; the wings hyaline, iridescent, with the veins brown;
front wings with the radio-medial cross-vein present; the free part of
M,+Cu, received near the middle of the cell M,; the stigma about three
times as long as wide; hind wings with the cell R,+, broadly open at
apex; the saw-guides narrow, the upper margin straight, apex rounded,
curved from the upper apex to the lower base. Length 4 mm.
Habitat.—Canada (Marlatt). Types in the collections of
the American Entomological Society and of the U. S. National
Museum.
Described from notes made by Mr. 8S. A. Rohwer for the
writer from the type in the collections of the American Entomo-
logical Society.
Metallus capitalis Nort.
Body reddish-yellow with the legs pallid and the head, the anten-
nae beyond the second segment, the mesopleurae, the pectus, and the
apex of the abdomen, piceous; the head with an indistinct transverse
carina between the antennae; the antennal furrows distinct to about
half the distance to the median ocellus; the median fovea present, the
lower wall more distinct; the ocellar basin with the upper lateral walls
not strong; the interocellar furrow not strongly continuous; the an-
tennae rather long and slender, rather hairy, the third segment a little
1909 | American Species of Scoltoneurinae 267
longer than the fourth and not as long as the fourth and fifth together,
the eighth and ninth subequal; the wings hyaline, the venation pale
brown; the front wings with the radio-medial cross-vein represented
only by minute rudiments; the free part of M,+Cu, received in the
cell M, distinctly beyond its middle; the stigma twice as long as wide,
angled at middle and at the point of attachment of the radial cross-
vein; the hind wings with the cell R,+, closed and appendiculate at
apex; the saw-guides straight above, obliquely truncate, and rounded
below. Length 3.5 mm.
Habitat.—Brooklyn, New York. Type in the collections of
the American Entomological Society.
Described from notes made by Mr. 8S. A. Rohwer for the
writer from the type.
The present synopsis was prepared for publication several
months ago, it was not sent off because of the uncertainty felt as
to the status of the species canadensis, capitalis, and rubi. My
views regarding the validity of these species based on the data
and specimens at hand were written up ready for publication
and the description and notes were loaned to Prof. C. O.
Houghton* and included by him in an article on ‘“‘The Black-
berry Leaf-Miner.’’ The description there given is based in
great part on the specimen described in this paper as Metallus
rohwert. The careful study made of the types of canadensis
and capitalis by Mr. Rohwer, together with the accumulation of
several additional specimens has convinced me that I was in
error and that all these forms should be ranked as distinct species.
Metallus rohweri n. sp.
Body black with the two basal segments of the antennae more or
less, the middle and lateral lobes of the mesonotum, the scutellum,
and its appendage, and the postscutellum, rufous; the legs with the
trochanters, the knees, the front and middle tibiae and tarsi, the post-
terior tibiae, more or less infuscated toward the apex, and the posterior
tarsi yellowish-white; the antennal furrows extending a short distance
from the antennal fovea, interrupted and forming a deep well-like
puncture, interrupted on the middle of the front to opposite the
median ocellus, continued around the outer side of the lateral ocellus
to the occiput, the vertical portion of the antennal furrows punctiform,
longer than wide; the interocellar furrow deep and straight; the post-
ocular area strongly convex and distinct; the ocellar basin elevated,
flat, forming an indistinct pentagonal area, depressed in front of the
median ocellus and a square depressed area behind connecting with
the interocellar furrow; the supraclypeal area not strongly elevated,
uniformly convex; the median fovea broad, deep, and prominent,
* Houghton, C. O.—Entomological News, XIX, 1908, 214.
268 Annals Entomological Society of America [Mol is
slightly papillate at bottom, situated between and above the bases of
the antennae; the antennae long and slender, the antennae with the
first and second segments more or less annular, broader than long, sub-
equal in length, much shorter than the third, the third longer than
the fourth and shorter than the fourth and fifth together, the fifth and
following segments subequal, the ninth segment broadly and rather
sharply pointed; the wings, including the veins, the stigma, and the
costa, brownish; the front wings with the radio-medial cross-vein
present, hyaline; the free part of R, about twice the length of R,;
the free part of M,+Cu, received in the middle of the cell M,; the stig-
ma about twice as long as broad, obliquely truncate beyond the radial
cross-vein and uniformly convex before it; the hind wings with the
cell R,,, closed and appendiculate; the posterior metatarsus slightly
shorter than all the following segments together, the second segment
one-half the length of the metatarsus and subequeal in length to the
third; the saw-guides straight above, convex below, obliquely trun-
cated at apex, rounded at angle below, and sharply pointed at upper
apical angle. Length 4.5 mm.
Habitat.—Block Island, Rhode Island (A. P. Morse, Col-
lector) ; Missouri (?), larva on blackberry, Riley Collection.
Since the above was written two additional specimens have
been received from Dr. F. H. Chittenden, bred by F. C. Pratt,
from a leaf-miner collected on blackberry at Colmanville, Penn-
sylvania. These specimens are identical with the type specimen
described above and one of them has been labelled as a paratype.
This species is named for Mr. 8. A. Rohwer, to whom I am
indebted for the notes on the described species of Metallus given
above.
Metallus rubi Forbes.
Body black with the trochanters, the knees, the front and middle
tibiae and tarsi, the hind tibiae, though more or less infuscated, and
the hind tarsi, white; the body color varies from brownish to jet black;
the antennal furrows distinct above the base of the antennae, puncti-
form, varying from a deep well-like puncture to one with flaring sides,
some individuals show another puncture on the margin of the antennal
foveae, connected with it, and sometimes connected with the above
described puncture above the bases of the antennae; antennal furrows
obsolete on the front, line-like impressions behind the lateral ocell,
faint along their outer margin and distinct and finely punctiform in
front of them; postocular area strongly convex, well separated; the
interocellar furrow distinct, concave behind; the ocellar basin elevated,
indicating a pentagonal area, flat or slightly convex on the outer sur-
face, depressed close to the median ocellus, the depression connecting
with the interocellar furrow; the median fovea situated between and
above the bases of the antennae, circular, sometimes small and well-
like, sometimes broad with flaring sides; the supraclypeal area uniform-
1909] American Species of Scolioneurinae 269
ly convex; antennae long and slender, the first and second segments
together one-half the length of the third, the third longer than the
fourth and shorter than the fourth and fifth together, the fifth and fol-
lowing segments subequal in length, the ninth segment strongly con-
vex on the basal third and gradually, convexly, tapering to a point,
the third and following segments more or less serrate on one side; the
front wings with the radio-medial cross-vein present, hyaline, or want-
ing; the free part of M,+Cu, received near the middle of the cell M,;
the stigma twice as long as wide, broadly convex on the lower margin
and at apex; the free part of R, twice the length of the free part of R,;
the hind wings with the cell R,;, closed and appendiculate at apex;
the wings varying from brownish to hyaline, the veins, stigma, and
costa black or brown; the posterior metatarsus distinctly shorter than
all the following segments together, the second segment longer than
the third and almost half the length of the metatarsus; the saw-guides
broad, straight above, convex below, obliquely truncate at apex,
broadly rounded at lower angle, and rather sharply pointed at upper
apical angle. Length 3-4 mm.
Habitat.—Ilinois (Forbes); Dover, Delaware (Houghton);
New York (Cornell Agricultural Experiment Station); River
de Pere and Central Missouri, Riley Collection. Bred from
larvae found on cultivated blackberry and on dewberry.
An extremely variable species in coloration and head structure.
This may represent more than one species but the material at
hand does not warrant its subdivision at the present time. The
male differs only in its smaller size and in having the antennae
more decidedly, flattened and saw-toothed than in the female.
Entodecta Knw.
Head transverse, the front not as broad as high; the inner
margin of the eyes uniformly convex, the eyes distinctly con-
verging below; the antennal furrows obsolete except adjacent to
the antennal foveae and behind the lateral ocelli; the clypeus
slightly emarginate, not more than half as wide at apex as at
base; the antennae long and slender, filiform, the second seg-
ment distinctly shorter than the first, wider than long, annular,
not so wide as the first, the first and second together distinctly
shorter than the third; the front wings with the free part of R,
and the radial cross-vein not interstitial and inclined at the same
angle; the free part of M,+Cu, received distinctly before the
middle of the cell rst A+2nd 2nd A; media gently curved at
the point of separation of M,,, and M,,,; the radial sector almost
straight, oblique at base. Type Tenthredo pumila Klg.
to
70 Annals Entomological Society of America [Vole aii:
The generic diagnosis given above is based upon a speci-
men of the type species purchased from Dr. O. Staudinger and
A. Bang-Haas. This diagnosis is entirely different from that
given by Konow. He separated the genera Scolioneura and
Entodecta on the position of the median ocellus, in the first genus
this ocellus is said to be below a line drawn between the dorsal
margins of the eyes and above in the later. Konow says of
Entodecta: ‘‘ Oeberkopf hoch tuber die Augentangenten empor-
steigend; das unter Nebenaugen liegt noch tiber dieser Tangente ”’
and of Scolioneura: ‘‘ antere Ocelle unter den Augentangente.”’
An examination of Eutodecta pumila and Entodecta get, the latter
specimen received from Konow, shows this ocellus located dis-
tinctly below and not above the “ Augentangente;”’ in fact it
occupies the same position in all the species that I] have examined
and the character is therefore entirely worthless for the differen-
tiation of these genera. The head is slightly more elevated behind
the median ocellus in Entodecta pumila but not as wide as in
Scolioneura betulett.
The specimen of Entodecta alaskana Kincaid was not avail-
able for examination when the above description was prepared.
Entodecta humilis Knw. is unknown to me and a copy of his des-
cription is given below. These species may not belong to Ento-
decta as here defined.
Species of Entodecta.
a. Abdomen black; the tegulae and the legs beyond the middle of the femora
DrOwilish-whites vec se tine ee oe eee eater alaskana Kced.
aa. Abdomen fuscous; the tegulae black; the legs yellowish white. . humilis Knw.
Entodecta alaskana Kcd.
Body black with the tegulae, the costa, the stigma, the wing veins,
and the legs below the middle of the femora, brownish-white; the an-
tennal furrows continuing as a shallow furrow from the cylpeal fovea
to the occiput, expanding above the bases of the antennae into a pit
twice as long as broad, ending in a circular pit behind the lateral ocelli,
extending in front of the lateral ocelli as a deep furrow about the width
of an ocellus; the postocullar area short, the interocellar furrow shal-
low; the ocellar basin a V-shaped depressed area around the median
ocellus, joining the interocellar furrow behind; the cheeks convex, not
at all declivous; the malar space usually broad; the supraclypeal area
prominent, strongly arched; the median fovea inconspicuous, elongate,
about one-fourth the width of the space between the antennae; the
antennae with the second segment elongated, about one-half the length
of the first, the third segment as long as the fourth and fifth together;
1909] ' American Species of Scolioneurinae 270
the posterior metatarsus as long as the two following segments togeth-
er; the front wings with the radial cross-vein almost interstitial with the
free part of R,. Length 4 mm.
Habitat.—Kukak Bay, Alaska. Description drawn from
Kincaid’s unique type.
Entodecta humilis Knw.
‘Niger; abdomine fusco; labro, palpis, pedibus flavescenti-albidis ;
unguiculis subfuscis; antennarum 2 articulis basalibus e fusco lutes-
centibus; alis fuscenti-hyalines; venis et stigmate fuscis.
Parvus, nitidus; capite et thorace pilis tenuibus, brevibus, sparsis
obsitis; capite pone oculos fortiter angustato; clypeo apice truncato;
antennis gracilibus, compressiusculis, breviter cano-pilosulis, trun-
cum longitudine fere aequantibus; fronte puncto impresso supra-
antennali ornata; area frontali deleta; vertice brevi, longitudine sua
duplo latiore. Lng. 3.5 mm.
RPatia: Alaska (Sitcha):
Von Herrn F. Sahlberg gesammelt. Die Art unterscheidet
sich von den verwandten durch den nicht ausgerandeten sondern
gerade abgestutzten Clypeus sowie durch die kleine tiefe punkt-
formige Supraantennalgrube, und ist unter den bisher bekannten
4 Arten der Gattung die kleinste.”’
LIFE HISTORY OF CORIZUS LATERALIS SAY.*
By J. C. HAMBLETON.
This insect was first described by Say 1n 1825 under the name
Coreus lateralis. Again in 1852 Dallas described it as Rhopalus
punctipennts. Later, in 1859, Signoret in his ‘“‘Monograph of
the Genus Corizus’”’ acknowledged both species, having been
guided largely by color characters.
There are in all twelve known species of the genus in North
America,f this being one of the most widely distributed, though
apparently confined to the United States. It has always been
a source of trouble to collectors because of its extreme variability
in color.
Up to the present, little has been known and nothing pub-
lished in regard to the life history or habits of any of them.
Prof. Herbert Osborn found nymphal forms of hyalinus Fab. a
few years since on a species of Euphorbia, but has published no
complete description of its stages.
Lateralis was found this past summer in great abundance in
the neighborhood of Columbus, Ohio, wherever its host plant was
found. This is Polygonum pennsylvanicum a native annual that
is quite generally distributed over the whole of the United States.
Adult insects could always be taken also on colonies of Poly-
gonum persicaria, a species introduced from Europe, and also
widely distributed, however neither eggs nor young were found
upon it. Several broods of young were reared to maturity on
persicaria in the laboratory and no difference could be observed
between them and those reared on pennsylvanicum where
the eggs were deposited. The adult forms fed freely upon perst-
carta in captivity and apparently do so in the field since they could
always be taken on colonies of this plant, but as aforesaid, dili-
gent search failed to reveal either eggs or young. This is evident-
ly a case of partial adaptation that has taken place since the ad-
vent of the plant to this country. The question may be asked,
why does the female not deposit her eggs on this plant as well
as on the other when she herself feeds upon it and when her
young will thrive upon it?
* Read at the meeting of the Ohio Academy of Science, at Delaware, O.,
Noy. 26th. 1909.
7 Ann. Ent. Soc. of America, Vol. I, p. 133.
{ Bull. No. 46, U. S. Dept. of Ag. Div. of Entom. p. 89.
272
1909] Life History of Corizus lateralis Say. 278
A close examination of the two plants in question will, at least,
suggest an answer. The petioles of pennsylvanicum, where the
eges are deposited, are thickly beset with glandular hairs, while
those of persicaria are entirely smooth. These hairs may act as
the immediate stimulus to oviposition and also may serve the
insect in a mechanical way during the act.
Eggs were first found early in August, and at this time young
in all stages of development were also found so that oviposition
must have begun early in July, and perhaps earlier. The eggs
are usually deposited on the petioles in small clusters of from
two to twelve, a few centimeters below where the flowers begin,
though occasionally they can be found singly and also sometimes
among the blossoms themselves. They are not attached directly
to the plant, but by means of a fine thread from one to two mili-
meters long. They are at first of a light yellow waxy color.
This changes to a bright red as the time for hatching approaches.
This occurs apparently in about six days, though this could
not be definitely determined as no female was observed depositing
her eggs, and none did so in captivity. Six days was the longest
time any eggs delayed in hatching after being collected.
It seems probable that more than one brood may be reared
during the summer. Either this is the case or the egg-laying
period is very much prolonged, for eggs and young were found
until the autumn frosts had killed the host plants.
The young when first hatched, hike each instar when first
moulted, is quite red in color. Upon drying and hardening the
head and thorax turn black and the abdomen a light green.
After feeding, the latter becomes a pale greenish yellow. These
colors are kept unchanged until the first moult though toward
the latter part of the first instar a faint red spot is seen just
above the dorsal glands, and the black of head and thorax shows
a reddish tinge. Before feeding, the rostrum which is carried
beneath the body is longer than the insect itself and can be seen
protruding beyond the extremity of the abdomen.
Within two hours after hatching, the young find their way up
the stem to the blossoms. The petals of this plant are persistent
and enclose the young seed until it is ripe.
The rostrum is inserted into these and search is apparently
made for the young seed, for the rostrum can be seen to be thrust
in at different angles without taking it out of the first incision.
When they have found the spot that suits them the abdomen
to
74 Annals Entomological Society of America [Molain:
soon becomes distended and is henceforth quite the most promi-
nent part of the insect. They remain during the whole of this
instar hidden among the blossoms. Their small size permits
them to do this with ease. Their color during this stage is decid-
edly protective, as indeed it is during all stages. The color of the
petals is pink and red while the stem and lower parts of the blos-
soms are a pale green. The black of the head and thorax blends
perfectly with the shadows while the paler color of the abdomen
harmonizes with that of the stems.
This instar lasts from two to threedays when the insect anchors’
itself by extending its fore legs as far forward as possible and its
hind ones far back. The skin splits open longitudinally on the
thorax and the insect slowly pulls itself out. At first it is a pale
red, but this deepens and in some cases passes to a decided brown.
The color of the entire insect is now quite uniform except on very
close inspection. The head and thorax though slightly darker
than the abdomen have permanently lost their black color.
This is in keeping with their new surroundings for their increased
size no longer permits them to remain below in the shadows but
they must now remain outside where the prevailing colors are |
red and pink. Toward the latter part of this stage the wing-pads
may be faintly distinguished beneath the skin, and two black spots
appear on the abdomen, one on either side of the dorsal glands.
This instar lasts from two to four days.
The most notable change in the third instar is the appearance
of the wing-pads clearly outlined, with their veins showing as
black stripes.. The thorax is also more clearly defined and the
eyes have become more prominent. This instar is of from three
to five days duration.
In the fourth stage, which lasts from five to seven days, the
head is clearly defined by the constriction between it and the
thorax. It now has more nearly the form of the head of the adult
insect. The prothorax has also become more prominent, and the
scutellum becomes faintly visible. The wing-pads have become
quite prominent with their outer margins and three veins black.
The fifth and last instar lasts from three to nine days. The
scutellum, in this stage, is clearly defined and the wing-pads
have become very prominent, reaching almost to the fourth abdo-
minal segment. The antennae have apparently reached their
full development though the tarsi are, as in all the nymphal
stages, but two jointed. The eyes are very well developed but
i
1909] Life History of Corizus lateralis Say. as
the ocelli have not yet appeared. The body is now almost as
large as that of the adult and has the same general appearance.
There is one character common to all the nymphal stages of
this insect except the first, which gives some insight into its
phylogeny. This is the banded legs. These are lost in the adult
lateralis, but are present in sidae, its tropical relative, which
barely reaches our southern borders. This seems to indicate that
probably sidae is the progenitor of lateralis and that this change
has been the effect of climate.
While the different stages may vary much in duration that
of the sum total of all does not. This is, in all cases observed,
from twenty to twenty-one days.
There yet remains one important fact to be learned in the
life history of this insect. It is not known positively how it
passes the winter. There can be little doubt, however, that, it
hibernates in the adult stage. It is inconceivable that either
the eggs or young could endure the winter after their host plant
has perished. As before stated, young could be found in all
stages until frost, and then all, including the adults disappeared.
It is probable that the former were killed while the latter sought
shelter among rubbish.
It was not definitely determined whether the host plant 1S
injured by the attack of this insect or not. There was, however,
some indication that such is the case. One colony that harbored
an extraordinary number of insects ripened very few seed.
Some heads not having more than a half dozen seed that reached
maturity.
Many of the eggs collected failed to hatch. Instead of turn-
ing red they soon became almost black and quite opaque. Later,
from these eggs small hymenopterous parasites would emerge,
whose identity has not yet been determined.
DESCRIPTION.
EGG.
In size, .5 by .7 mm.,—oval when viewed from the side but
somewhat triangular when viewed from the end.
Color: Yellowish, pale, turning red before hatching. Attached
by slender thread to petioles of host plant in clusters of two to twelve.
FIRST INSTAR.
1 mm. in length. Head and thorax prominent, not separated
by constriction. Abdomen small, consisting of nine segments, the
fifth of which is interrupted at the middle and two small glands are
276 Annals Entomological Society of America [Vol. IT,
placed between. Antennae of four segments, as in the adult. Legs
similar to those of adult except the tarsi which are two-jointed.
Color: Head and thorax black, abdomen greenish, changing to
yellowish green. Antennae and legs reddish. Whole insect covered
with stiff hairs which are prominent on the head.
SECOND INSTAR.
2mm. in length. Head and thorax as in first instar, except as
to color.
Color: Whole insect of a reddish cast due to small red spots and
blotches on a yellowish background. Antennae with two narrow dark
stripes extending through first three segments. 4th segment darker
toward the vertex. Two black spots on interrupted segment, one on
either side of dorsal glands. Legs yellowish banded with red. Entire
insect with coarse hair.
THIRD INSTAR.
3mm.inlength. Eyesimore prominent, Wing-pads appearing
with veins black. Two black spots on 2nd abdominal segment, 2
on 4th and 2 on 5th. Other characters as in second instar.
FOURTH INSTAR.
4 mm. in length. Head separated from thorax by constriction.
Prothorax clearly defined. Wing-pads more prominent. Abdomi-
nal spots sometimes more numerous, though the six of the third instar
always present. Scutellum faintly outlined by lighter shade.
FIFTH INSTAR.
5mm. in length. Wing-pads reaching almost to 4th segment of
abdomen, much darker in color especially toward the extremities.
Spots on 2nd segment increased to four, others as above.
DESCRIPTION OF PLATE.
Cluster of eggs attached to host-plant.
First instar of Corizus lateralis Say.
Second instar of Corizus lateralis Say.
Third instar of Corizus lateralis Say.
Fourth instar of Corizus lateralis Say.
Fifth instar of Corizus lateralis Say.
~~
ee
a
7)
ANNAIS HK. S. A. VoL. II, PLATE XXIX.
J. C. Hambleton.
INDEX TO VOLUME II.
Achaetoneura, 249.
Acraeinae, 223.
Adaptation palpus of male spiders, 60.
Adelpha,: 230, 231.
Agalenidae, 63.
Agropyron
occidentalis, 72.
tenerum, 72.
Aleurodes, 121.
Aleurodidae, 101, 121.
Aleurodiscus, 122.
Allobia, 93.
Alopecurus geniculatus, 72.
Andrews, title of article, 3.
Anisomorpha buprestoides—
host of Phasmophaga, 244.
Anomoneura mori, 120.
Anosia, 209, 210.
plexippus, 209, 211.
Anosia scent pockets of, 3.
Antiphrisson, 140.
Apatura, 221.
Aphelinus nigritus, 77.
Aphididae, 30, 101.
Aphididae, 196.
Aphis atriplicis, 88, 89, 92, 93, 94, 95.
betulaecolens, 30.
brassicae, 85, 86.
carbocolor, 93.
cephalanti, 40.
genistae, 95.
gossypu, 86.
graminum, 68.
maidiradicis, 86, 938.
maidis, 86.
mali, 93.
nymphee, 95.
rumicis, 89, 98.
sambuci, 95.
setariae, 86.
tarmentille, 95.
Aphycus, 93.
Araschnia levana, 221.
Argiopidae, 63.
Argynnis, 229.
diana, 229, 230.
niphe, 212.
Aristolochia, 222.
Asaphes rufipes, 93.
Asilus, 136, 140.
affinis, 140, 142, 153.
albicomus, 141, 1438, 150.
angustipennis, 140, 143, 152.
antimachus, 141, 158.
arizonensis, 140, 142, 167.
astutus, 140, 141, 163.
avidus, 140, 144, 147.
auriannulatus, 140, 142, 151.
279
auricomus, 141, 143, 148.
brevicomus, 140, 143, 155.
cacopilogus, 140, 141, 165.
californicus, 141, 143, 164.
callidus, 141, 144, 157.
coquilletti, 140, 143, 154.
delusus, 141, 144, 159.
erythocnemius, 141, 144, 163.
flavofemoratus, 140, 142, 153.
fuscatus, 140, 142, 168.
gracilis, 141, 142, 145.
johnsoni, 141, 144, 159.
latipennis, 140, 143, 152.
lecythus, 140, 144, 145.
lepidus, 141, 143, 150.
leucopogon, 140, 141, 166.
limpidipennis, 140, 142, 167.
maneei, 141, 144, 158.
mesae, 141, 1438, 162.
midas, 140, 141, 165.
montanus, 141, 143, 149.
nitidifacies, 140, 141, 165.
notatus, 141, 144, 157.
novae-scotiae, 141, 144, 157.
occidentalis, 140, 142, 147.
orphne, 140, 142, 154.
paropus, 141, 144, 161.
piceus, 141, 143, 149.
prairiensis, 141, 144, 161.
rubicundus, 141, 144, 162.
rubidus, 144—rubicundus, 144.
rufipennis, 140, 142, 168.
sadyates, 141, 144, 156.
sericeus, 140, 141, 164.
snowi, 141, 144, 160.
terminalis, 140, 142, 155.
tenebrosus, 140, 144, 146.
truquii, 140, 142, 169.
tuxpanganus, 140, 142, 169.
willistoni, 141, 1438, 150.
Atriplex patula, 88.
Baccha dimidiata, 254.
Basilarchia, 209, 232.
Baumhaueria, 245.
Beta, 88.
Betula, 260.
Bibionid, 3.
Bigonichaeta, 245.
Birch Callipterus, 30.
Blennocampa, 259.
Brachychaeta, 245.
Brachyopa notata, 254.
Bromus secalinus, 73.
Brues, C. T., title of article, 3.
Butterflies, Mimicry in N. A., 203.
Button bush aphid, 40.
280
Calaphis betulella, 30.
‘allipterus, 30, 33.
betulae, 30.
betulaecolens, 30.
tiliae, 33.
ulmifoli, 115.
Caudell, A. N., title of article 3.
Camptotera, 3.
Camptotera catalogue, 22.
papaveris, 22, 24, 25.
pulla, 23, 27.
clavata, 23, 26.
metotarsa, 23, 26.
Cecidomyiidae, 3.
Cephalanthus occidentalis, 40.
Cerdistus, 141.
Cethosia, 212.
Chaitophorus, 86.
populicola, 116.
Chalcidoma muraria, habits of, 9.
Chalcis flies, seed infesting, 3.
Chamberlin, R. V., article by, 175.
Chamberlin, R. V., title of article by, 3.
Chenopodium, 88, 90, 91, 98.
Chermes, 93. 113.
Chermes abietis, 110, 112.
pinifoliae, 111.
Chlorippe, 221.
Clubionidae, 63.
Coccidae, 101, 122.
Cockerell, T. D. A., article by, 251.
Comstock, J. H., article by, 9.
Comstock, title of paper,
Coquillett, D. W., title of paper,
Corizus, 272.
hyalinus, 272.
lateralis, 272.
Corydalis cornuta, 3.
tracheal supply of nervous system of
larva, 46.
Cordyligaster, 250.
septentrionalis. n. s., 250.
Crosby, C. R., title of paper, 3.
Cyclotaphrys, n. g., 246.
anser n. s., 246.
Dacytlis glomerata, 72,
Danaida, 211, 213, 222
berenice, 211, 213, :
chrysippus, 211, 21:
decipiens, 212.
genutia, 211, 212, 213.
insolata, 212.
plexippus, 211, 213, 2
220) 222:
Danainae, 223.
Danaini, 208.
Dasyllis fernaldi, 256.
Dasypogon, 255, 256
teutonus, 2:
Index to Volume II
Davis, J. J., artieles by, 3750; 197.
Death feigning in Zaitha, 3.
Diapheromerata femorata—
host of Phasmophaga, 244.
Dictynidae, 63.
Dioctria, 254.
florissantina, 254.
Diptera, fossil, 253.
Distichlis spicata, 72.
Drepanosiphum, 196.
acerifoli, 196.
braggi, 196.
platanoides, 196.
Dysdera crocata, 62.
Dysderidae, 62.
Eleusine indica, 73.
Elymniinae, 212.
Elymus canadensis, 73.
Strata, (2:
Entodecta, 259, 261, 270.
alaskana, 270.
humilis, 270, 271.
pumila, 270,
gei, 2f0:
Entomophora aphidis, 89.
Epitriptus, 141.
Eragrostis, pilosa, 73.
megastachya, 73.
Eriocampoides, 252.
revelatus n.s., 251.
Erycinidae, 221.
Escaryus phyllophilus, 177.
Eucallipterus tillae, 33.
Eucelatoria, n. g., 249.
armigera, 249.
Eumasicera, n. g., 248, 249.
coccidella, n. s., 249.
Eumenidae, 252.
Euonymus atropurpurea, 98.
Eupeodes, 253.
Eupitenus rediscovery, 3.
Euploea, 211.
brenchleyi, 212.
asyllus, 212.
Euploeini, 208.
Euproctis chrysorrhoea,
host of Cyclotaphrys, 246.
Euryceromyia, 245.
Euthyprosopa, 244, 245.
Eutolmus, 140.
Eutresis imitatrix, 208.
Evolution palpus of male spiders, 60.
Executive Committee, Report of, 3
Exorista griseomicans, 248.
Felt, E. P., title of paper, 3.
Fenusa, 259. .
Figites, 93.
Index to Volume II
Filistatidae, 62.
Forbes, S. A., title of article, 3.
Formica fusca, 38.
Fossil, Insects, 251.
Frontina, 249.
Gall aphid, 88.
Gall insects, nomenclature of, 8.
Geophilidae, 175.
Geophilus atopleurus, 181.
attenuatus-unaster, 179.
bipuncticeps, 178.
deducens, 180.
claremontus, 184.
glaber, 185.
isantus, 185.
legiferens, 182.
marginalis, 183.
nasintus, 183.
occidentalis, 185.
regnans, 185.
rubens, 182.
smithi, 182.
Geum, 260.
Girault, A. A., title of article, 3.
Gnaphosidae, 63.
Grapta, 221.
Hairs of spiders, 14.
Hamadryas, 208.
Hamamelistes, 113.
spinosa 93, 114.
Hambletons J. ©. article by, 272.
Haplophilus audacior, 177.
laticeps, 177.
teniopsis, 177.
Hayhurst, Paul, article, by 88.
Hayhurstia, 94, 95.
Heliconiinae, 208.
Heligmoneira, 140.
Hemerocampa leucostigma,
host of Sisyropa, 248.
Hestia, 208..
Hilton, W. A., article by, 46.
title of paper, 3.
Hippodamia 13-punctata, 94.
Homologies of wing. veins, 101.
Hooker, W. A., title of paper by, 2.
Hordeum caespitosum, 72.
pusillum, 72.
Hormaphis hamamelidis, 93.
Hymenoptera, fossil, 251.
Hypertrophocera, 244, 245.
Hypolimnas missippus, 212, 213.
Idiopterus n. g., 198.
nephrelepidis, 199.
Ithomiinae, 208.
Ituma, 208.
phenarete, 208.
to
oe)
ei
Lachnus nudus, 93.
piceicola, 93.
Laphria fernaldi, 256.
Lasius niger, 38.
Limbas, 210, 211.
chrysippus, 210, 211, 212.
var dorippus, 210.
Limenitis, 208, 214, 220, 221, 222, 231
234.
archippus, 209, 214, 232.
anthems) 24 215, 26s. 219) 227
228, 230, 232.
astyanax, 219, 227, 228, 229, 230.
bredowi, 231, 233, 234.
californica, 230, 231, 232, 233, 234,
DOP
eros, 217.
floridensis, 217, 218, 219, 229.
hulstie2ilG, 218% 21/9:
proserpina, 227.
lorquini, 230, 231, 232, 234, 235.
sybilla, 214, 215.
wiedermeyeri, 233.
Linden Callipterus, 33.
Linotenia bidens, 186.
chionophila, 186.
fulva, 186.
lebipes, 186.
rubelliana, 187.
Lithobiidae, 175.
Lithobius aztecus, 191.
bipunctata, 191.
bonvillensis, 189.
eeler,, 190:
clavigerus, 187.
forficatus, 190.
fulvicornis, 191.
ginampus, 187.
harrietae, 191.
hoples, 191.
kochii, 187.
monticola, 190.
mordax, 190,
multidentatus, 190.
obesus, 187.
paradoxus, 187.
pusio, 187.
paucidens, 191.
sierravagus, 190.
sokovus, 189.
tivius, 188.
utahensis, 187.
xanti, 190.
Liriodendron tulipifera, 36.
Lycorea, 208.
Lycoreini, 208.
Lycosidae, 63.
Lysiphlebus, 69, 79, 80, 81, 85, 86.
tritier, (i, te:
eragrostaphidis, 93.
282 Index to Volume II
Machimus, 140.
Macrosiphum cynobasti, 38.
liriodendri, var. rufa, 36.
pisi, 114.
solanifolii, 117.
Masicera, 249.
armigera, 249.
McGillivray, A. D., article by, 259.
Megachile praedicta, : 253.
Megaxyela petrefacta, 252.
Melanophrys, 245.
Melanoxanthus salicis, 93.
Melospiza melodia, 77.
Metallus, 259, 261, 265.
canadensis, 266, 267.
capitalis, 266, 267.
rohweri, 266, 267.
rubi, 259, 260, 265, 266, 267, 268.
Microstylum, 254.
destructum, 254.
Mimetidae, 63.
Mimicry in N. A. Butterflies, 203.
Mindarus, 114, 115.
Muscular system spiders, 16.
Myzuz cerasi, 110.
Nelson, J. A., title of paper, 3.
Neoitamus, 140.
Neotractocera, 245.
Nicocles, 255, 256.
miocenicus, 255.
TULUS LOO 200.
Nigrasilus, 140.
Nomenclature, Report of Committee, 8
Odynerus terryi, 252.
Officers, 2.
O’Kane, W. C., article title, 2.
Ooctonus, 22.
Oxyopidae, 63.
Pachyneuron micans, 93.
Pachypsylla, 120.
c-mamma, 120.
Palpus of male spiders, 60.
Papilio, 222, 223, 224, 230.
antennor, 222.
asterius, 225, 226, 227, 228.
glaucus, 227.
glaucus glaucus, 228.
glaucus canadensis, 225
machaon, 222.
melasina, 226.
polyxenes, 225.
polyxenes asterius, 225
polyxenes americus, 226,
philenor, 226.
troilus, 226.
troilus troilus, 225.
turnus, 225, 227.
Parachaeta, 250.
bicolor, 250.
inermis, 250.
Parabates, 261, 262.
histrionicus, n. s., 262, 263.
inspiratus n. s., 263, 264.
Paragermania .o-.22e/e
autumnalis n. s., 247.
Patch, Edith M., article by, 101.
Pectiniunguis heathu, 176.
monterus, 176.
Pemphigus, 113.
acerifolii, 117.
Pemphigus attenuatus, 93.
Petrunkevitch, A., article by, 11.
title of article 3.
Pharmacophagus, 222, 223.
hahneli, 223.
philenor, 224, 228, 229, 230.
polydames, 924,
Phasmophaga n. g., 243, 244, 245
antennalis, 243, 244.
meridionalis, 244.
Pholcus phalangoides, 14.
Phoridae, 3.
Phylloxera, 93.
Phymatosiphum, 196.
monelli, 197.
Pisauridae, 63.
Poa pratensis, 72.
Polybates, 261.
Polybates, 261, 264.
slossonae, 264, 265.
Polyganum pennsylvanicum, 272.
persicaria, 272.
Polygonia, 221.
Porthetria, host of Tachina, 247.
Poulton, iD B., Address by, 203.
Proceedings, ik
Pseudatractocera, 245.
Pseudococcus citri, 123.
Pseudogermaria, 247.
Psylla floccosa, 117, 118, 119.
Psyllidae, 101, 117.
Pterocallis, 33.
juglandicola, 33.
Pteroclisis, 23.
Resolutions, Ashmead, 6.
Fletcher, 7
Ribes eynobastt 38.
Riley ella n. g., 249.
Riley, W. ie title of paper, 3.
Ritter, W. 1 title of article, 3.
Rhadiurgus, 140.
Rhopalus punctipennis 272.
Robberflies, 136.
Salatura, 210.
genutia, 211.
Index to Volume II
Scaeva, 253.
Scent pockets of Anosia, 3.
Schendyla nemorensis, 175.
Schizoneura, 113, 114.
rileyi, 114, 115.
Schizoneura americana, 93.
corni, 93.
lanigera, 93.
Schizoneura lanigera, 125.
Scolioneura, 259. 261.
Scolioneurinae, American, 259.
Scolioneurinae, 260.
Scolioneura, 270,
betuleti, 270.
capitalis, 260.
Scymnus americanus, 94.
Scytodes, 62.
Selandria capitalis, 259.
Severin, H. P., title of paper, 3.
Sherman, F. S., title of paper, 3.
Sicariidae, 62.
Siphocoryne, 94, 95.
Sisyropa, 248, 249.
hemerocampae, 248.
Skinner, Henry, title of paper, 3.
Smith, J. B., title of paper, 3.
Species, discussion of, 3.
Spiders, anatomy, etc., 11.
Spiders, evolution of palpus of male, 60
Spiders, palpi of, 3.
Sporobolus neglectus, 73.
Stilpnogaster, 140.
Stipa viridula, 72.
Syrphus, 253.
Tachina, 246, 247, 249.
larvarum, 246, 247.
japonica n. s., 247.
Tachinidae, new, 243.
Tasitia, 208, 210, 211.
berenice, 209, 210, 211.
berenice strigosa, 209.
Tetraneura, 113.
Thelymorpha, 245.
Theridiidae, 63.
Thomisidae, 63.
Thyridia, 208.
Ticks, host relation of, 2.
Tilia, 260.
Tolmerus, 141.
Townsend, article by, 243.
Toxoptera graminem, 2, 67, 69.
Tulip, tree aphid, 36.
Uloboridae, 63.
Uraphis, 94.
Urophylla, 245.
Wall Bee, habits of, 9.
Webster, F. M., article by, 67.
title of article, 2.
Wing viens, 101.
homologies of, 101.
Xylota pigra, 254.
Zaitha fluminea death feigning, 3.
283
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